Arthropods Vol. 8, No. 4, 1 December 2019 International Academy of Ecology and Environmental Sciences Arthropods ISSN 2224-4255 Volume 8, Number 4, 1 December 2019 Editor-in-Chief WenJun Zhang Sun Yat-sen University, China International Academy of Ecology and Environmental Sciences, Hong Kong E-mail: zhwj@mail.sysu.edu.cn, wjzhang@iaees.org Editorial Board Andre Bianconi (Sao Paulo State University (Unesp), Brazil) Anton Brancelj (National Institute of Biology, Slovenia) A. K. Dhawan (Punjab Agricultural University, India) John A. Fornshell (United States National Museum of Natural History, Smithsonian Institution, USA) Xin Li (Qingdao University, China) Oscar E. Liburd (University of Florida, USA) Ivana Karanovic (Hanyang University, Korea) Lev V. Nedorezov (Russian Academy of Sciences, Russia) Enoch A Osekre (KN University of Science and Technology, Ghana) Rajinder Peshin (Sher-e-Kashmir University of Agricultural Sciences and Technology of Jammu, India) Michael Stout (Louisiana State University Agricultural Center, USA) Eugeny S. Sugonyaev (Russian Academy of Sciences, Russia) Editorial Office: arthropods@iaees.org Publisher: International Academy of Ecology and Environmental Sciences Website: http://www.iaees.org/ E-mail: office@iaees.org Arthropods, 2019, 9(4): 110-117 Article Walking leg regeneration observed in three families and four species of antarctic sea spiders John A. Fornshell United States National Museum of Natural History, Department of Invertebrate Zoology, Smithsonian Institution, Washington DC, USA E-mail: johnfornshell@hotmail.com Received 1 September 2019; Accepted 27 September 2019; Published 1 December 2019 Abstract Observations of wound healing and regeneration of walking legs in specimens of Nymphon australe Hodgson, 1902, Nymphon charcoti Bouvier, 1911, Colssendeis tortipalpus Gordon, 1932 and Pentapycnon charcoti Bouvier 1910 archived in the U. S. National Museum of Natural History collections is reported. One hundred and ninety-four specimens of N. australe were analyzed for evidence of regeneration. Blastema formation and or regenerated limbs of reduced size were found in 64 individuals, 38%. Forty-four specimens of N. charcoti were analyzed for evidence of regeneration. A blastema and/or regenerated limbs of reduced size were found in 12 individuals, 27%. Fifteen specimens of Colssendeis tortipalpus were analyzed for evidence of regeneration. Five individuals, 33%, had either a fully regenerated walking leg of reduced size or a blastema was present. Ten individuals of Pentapycnon charcoti were analyzed for evidence of regeneration. None of these animals showed signs of regeneration. The Preferred Breaking Point (PBP), position of autotomy, was between coxa 1 and coxa 2. Regenerated limbs having all segments, but of reduced size were found in 5 specimens. In some cases, more than one walking legs had been regenerated or were in the process of regenerating as indicated by the presence of a blastema. A blastema formed on the end of the first segment of the chelophore was observed in a single specimen of N. australe. A blastema which was formed at the end of the second segment of the ovigerous appendage of C. tortipalpus was also observed. Keywords Nymphon australe; Nymphon charcoti; Colssendeis tortipalpus; Pentapycnon charcoti; regeneration; Preferred Breaking Point. Arthropods ISSN 2224­4255 URL: http://www.iaees.org/publications/journals/arthropods/online­version.asp RSS: http://www.iaees.org/publications/journals/arthropods/rss.xml E­mail: arthropods@iaees.org Editor­in­Chief: WenJun Zhang Publisher: International Academy of Ecology and Environmental Sciences 1 Introduction The members of the phylum Arthropoda are capable of limb regeneration including eye stalks in crustaceans after autotomizing the lost appendage (Bely and Nyberg, 2009; Flemming et al., 2007; Maruzzo, et al., 2005; Maruzzo et al., 2013). They cannot regenerate the whole body or the core or body axis (Bely and Nyberg, IAEES www.iaees.org Arthropods, 2019, 9(4): 110-117 111 2009). Regeneration has been described in members of the class Pycnogonida, order Pantopoda from the later part of the nineteenth century (Dohrn, 1881; Gaubert, 1892; Morgan, 1901; Loeb, 1905). The abnormal regeneration of the posterior portion of the trunk with a limb-like structure was described for Phoxichilidium femoratum (Rathke, 1799) [as Phoxichilidium maxillare Stimpson, 1853] by Loeb (1905). The regeneration of walking legs was reported by Dohrn (1881) and Gaubert (1892). More recent work has shown that immature stages of pycnogonids can regenerate their walking legs, but not mature adults that have lost the ability to molt (Bely and Nyberg, 2009). This regeneration includes the cuticle, digestive tract and in some cases the gonads (Fage, 1949; Hefler and Schottke, 1935). Other than review articles surveying the regenerative process in arthropods the subject has received little attention in recent years (Maruzzo et al., 2005; Maruzzo and Bortolin, 2013). Needham (1952) recognized two basic types of regeneration, epimorphosis and morphallaxis. Morphallaxis does not involve the proliferation of new cells; rather it results from the regeneration of the missing structures by the remodeling of existing cells. In morphallaxis, the remaining portion of the body is reorganized to restore the whole form. In epimorphosis, the former organ or limb is replaced by the direct development in situ of the portion of the animal which has been lost. Epimorphosis is typical of limb regeneration in arthropods. Epimorphacic regeneration requires cell proliferation and may be divided into nonblastemal and blastemal regeneration. Non-blastemal regeneration results from the trans-differentiation of the remaining tissue into the missing organ with limited proliferation of the surviving cells. Blastemal regeneration involves the formation of a specialized mass of cells known as a blastema. The latter is the type of regeneration found in the Arthropoda including the Pycnogonida (Mitić et al., 2010). In the process of epimorphic regeneration Needham (1952) identified two main phases, regressive and progressive. The regressive phase included: Wound closure; Demolition of damaged cells and defense against foreign organisms and chemicals and the dedifferentiation of cells to provide new tissues for the progressive or repair phase. The progressive phase is divided into the formation of the blastemal, the growth of the blastema or regeneration bud and the differentiation of the young regenerate (Needham, 1952). When a portion of a limb is lost or damaged it may be shed by the process of autotomy. In this process muscles constrict severing the portion of the limb distal to the Preferred Breaking Point (PBP) and sealing the end of the limb (Fleming et al., 2007). This process minimizes the damage to the remaining body of the animal and may also facilitate locomotion in the case of walking legs (Gross, 1969; Maginnis, 2006). In pycnogonids autotomy may also occur without subsequent regeneration as in the case of the shedding of cheliphores (Bain, 2003). The second and third larval appendages are lost in some species of pycnogonids only to be regenerated as palps and ovigerous appendages in the adults (Maruzzo and Bortolin, 2013). Regeneration in arthropods can only occur in conjunction with molting and is subject to endocrinological controls. These are natural results of the existence of the chitinous exoskeleton. Limbs may be autotomized at specific joints, a process which facilitates wound healing and future regeneration (Gross, 1969). The process of molting is stimulated to occur by the presence of ecdysones in the blood of the animal (Krishnakumaran and Schniderman, 1970). 2 Materials and Methods Archived specimens of Nymphon australe Hodgson, Nymphon charcoti Bouvier, Colssendeis tortipalpus and Pentapycnon charcoti Bouvier 1910 from the U. S. National Museum of Natural History were examined in this study for evidence of limb regeneration. Eight populations of Pycnogonida from the Antarctic Seas were analyzed in this study. Population A, USNM 1277471 including 29 specimens of N. australe. Population B including 77 specimens of N. australe collected at 760 0.2’ S. 1790 52.1’ W. Population C including 45 IAEES www.iaees.org 112 Arthropods, 2019, 9(4): 110-117 specimens of N. australe collected at 660 17’ 42’’ S. 1100 32’ 03’’E. Population D including 21 specimens of N. australe collected at 660 15’ 24’’ S. 1100 28’ 40’’ W. Population E including 23 specimens of N. australe, collected at 73o 49’ S. 1780 28’ 13’’ W. One population of N. charcoti including 44 specimens collected at 620 36.0’ S. 64014’30’’ W. One population of C. tortipalpus including 15 specimens, collected from 61o 18’ S. 56 o 09’ W to 610 20’ S 56 o 10’ W. including 10 specimens of P. charcoti each from a different location in the Antarctic seas. Because limbs of pycnogonids can become broken while being collected, preserved or subsequently being handled, a set of criteria was established for defining when regeneration had occurred or begun. The first step in regeneration is the formation of a blastema at the site of limb severance. Like many arthropods, pycnogonids have the ability to autotomize a damaged limb. This process of severing a limb in response to an external stimuli is a defense mechanism from attack by a predator or injury. This occurs at a joint between two segments termed the Preferred Breaking Point (PBP) (Fleming et al., 2007). Regenerated limbs are physically smaller, i.e. they are shorter and thinner than mature un-regenerated limbs on the same animal at the first molt following regeneration. The presence of such a reduced limb was taken as evidence of regeneration. Lost limbs, without the presence of a blastema were considered to be limbs lost in processing of the samples. These lost limbs were not counted as having been limbs in the process of regeneration. In some cases this may have resulted in an under counting of regeneration events. 3 Results 3.1 Nymphon australe Hodgson Population A: Ten adult and eight juveniles showed no signs of regeneration, limb loss or wound healing. One adult showed the results of regeneration producing smaller right walking legs 2 and 3 much thinner but the same length as normal walking legs. Two mature specimens showed early stages of regeneration in the form of a blastema on the 3rd and 4th right walking legs on one individual and a second individual with a blastema on the left 1st and 2nd walking legs. One specimen had blastemas on the 1st and 4th right walking legs. One specimen had blastemas on the 2nd and 3rd left walking legs plus the 4th right walking leg. One specimen had a blastema on the 1st and 2nd right walking legs. One specimen had blastemas on the 1st and 2nd left walking legs. One had blastemas on the 2nd and 4th left plus the 3rdright walking legs. One specimen had blastemas on the 2nd and 4th right walking legs. Two juveniles had lost limbs at the joint between the first and second coxa without a blastema, right 3rd walking leg and 4th left walking leg respectively. Unless otherwise noted the breaking point is between coxa 1 and coxa 2 (See Fig. 1). Population B: Thirty seven individuals or 48% had missing walking legs which had formed a blastema and two individuals with regenerated walking legs, one with the right second and third walking legs present in a reduced size and one individual with the first left walking leg and second right walking leg present in a reduced size (See Fig. 2). One juvenile had blastemas on the 2nd and 3rd right walking legs. One juvenile had a blastema on the 3rd left walking leg. Two juvenile specimens had blastema(s) on the 4th left walking leg. Three juveniles specimens had a blastema on the 3rd right walking leg. One juvenile specimen had blastemas on the 2nd, 3rd and left 4th right walking legs. One juvenile specimens had blastemas on the 3rd and left 4thright walking legs. IAEES www.iaees.org Arthropods, 2019, 9(4): 110-117 113 Fig. 1 The early stages of wound healing and regeneration of one of the walking legs of N. australe showing coca 1 and the blastema at the distal end of the autotomized limb. Fig. 2 A ventral view of N. australe with a regenerated left first walking leg. This limb is shorter and thinner than the other walking legs. IAEES www.iaees.org Arthropods, 2019, 9(4): 110-117 114 Population C: Twenty-seven adults and six juveniles showed no signs of regeneration, limb loss or wound healing. One juvenile had a regenerated left first walking leg of reduced size. One juvenile had lost the second and third segments of the left chelophore. A blastema had formed at the distal end of the basal segment. One juvenile had a blastema on the left ovigerous appendage. One juvenile had a blastema on the 1st right walking leg. One juvenile specimen had blastemas on the 2nd left and right walking legs. Seven juvenile specimens had blastemas on two or more walking legs. One juvenile specimen without wound healing had a break at coxa 1/coxa 2. Population D: Fifteen of the 21 adults showed no signs of limb regeneration. Six juveniles had initiated walking leg regeneration with the formation of a blastema. One specimen had a blastema on the 2nd right walking leg. One specimen had a blastema on the 2nd left walking leg. One specimen had blastemas on the 1st and 2nd right walking legs. One specimen had blastemas on the 3rd right and 4th left walking legs. One specimen had blastemas on the 1st2nd and 3rd walking legs. Population E: Thirteen adult animals showed no signs of regeneration. Ten juveniles had lost and regenerated one or more walking legs and/or initiated regeneration by the formation of a blastema. Two specimens had blastemas on the 3rd and 4th left walking leg. One specimen had a blastema on the 4thright walking leg and a fully regenerated 4th left walking leg. Two specimens had blastemas on the 4th left walking leg. One specimen had a blastema on the 2nd left walking leg and fully regenerated 1st and 4th right walking legs. One specimen had a blastema on the first segment of the left chelophore and a reduced size regenerated left first walking leg. One specimen had a blastema on the on the 3rd left walking leg. One specimen had a blastema on the 1st right walking leg. One specimen had a blastema on the on the 3rd right walking leg (See Table 1). Table 1 Summarizing the results for N. australe. The 193 missing walking legs all appeared to have been autotomized at the junction of coxa 1 and coxa 2. Left legs (I) (II) (III) (IV) Number of limbs with a blastema 25 26 26 25 Right Legs (I) (II) (III) (IV) Number of limbs with a blastema 16 17 34 24 3.2 Nymphon charcoti Bouvier 1911 Thirty-two of the 44 specimens, 76%, were adult animals showing no signs of regeneration. Two juvenile animals had fully regenerated but smaller walking legs one with the first right walking leg present as a smaller limb and one with the second left walking leg present as a smaller leg. Ten juvenile animals, 24%, had lost one or more walking legs and had formed a blastema as the first stage of regenerating the lost limb. One specimen had a blastema on the right 2nd and 3rd walking legs. One specimen had a blastema on the 3rd left walking leg. One specimen had a blastema on the 4th left walking. One specimen had a blastema on the 3rd right walking leg. One specimen had blastemas on the 2nd, 3rd and 4th right walking legs. One specimen had a blastema on the 3rd left walking leg. One specimen had a blastema on the right 3rd walking leg. One specimen had a blastema on the 4th left walking leg. One had a blastema on the 3rd left walking leg. One specimen had blastemas on the 3rd and 4th right walking legs (See Table 2). IAEES www.iaees.org Arthropods, 2019, 9(4): 110-117 115 3.3 Colssendeis tortipalpus Ten of the fifteen specimens of this species were adults and showed no signs of walking leg regeneration. One adult, however had lost the segments distal to the third segment of the right ovigerous appendage and a blastema had formed at the distal end of that segment (See Fig. 3). One of the juvenile specimens had a fully regenerated, but small, 4thleft walking leg. One juvenile had a blastema at the distal end of coxa 1 of the 1st right walking leg. Two juveniles had a blastema at the distal end of coxa 1 of their 4th left Walking leg. One juvenile had a blastema at the distal end of coxa 1 of the 1stleft Walking leg (See Table 3). 3.4 Pentapycnon charcoti Six juveniles and four adults were examined. None of these animals showed any signs of regeneration of limbs. These ten specimens were the total of all members of this species in the NMNH collections. Table 2 Summarizing the results for N. charcoti. The 9 missing walking legs all appeared to have been autotomized at the junction of coxa 1 and coxa 2. Left legs (I) (II) (III) (IV) Number of limbs with a blastema 1 1 2 1 Right Legs (I) (II) (III) (IV) Number of limbs with a blastema 0 1 2 2 Table 3 Summarizing the results for Colossendeis tortipalpus. The 5 missing walking legs all appeared to have been autotomized at the junction of coxa 1 and coxa 2. Left legs (I) (II) (III) (IV) Number of limbs with a blastema 1 0 0 2 Right Legs (I) (II) (III) (IV) Number of limbs with a blastema 1 0 0 1 Fig. 3 The left ovigerous appendage of Colossendeis tortipalpus with the blastema at the distal end of the third segment of the right ovigerous appendage. IAEES www.iaees.org 116 Arthropods, 2019, 9(4): 110-117 4 Discussion In his book, Pycnogonids, P. E. King (1973) states “If ambulatory limbs are broken off, the rupture usually occurs between the second and third segments of the limb in Nymphon gracile. Pycnogonids haveconsiderable powers of regeneration in the juvenile stages and when a break occurs the regenerative process is very rapid. After the stub of the ruptured limb is sealed off the new rudimentary limb is formed within the cuticle of the remaining part of the limb and is completed at a molt”. In this study the PBP in N. australe, N. charcoti, and C. tortipalpus is between the first and second coxa segments. The difference may simply be a species specific variation. In crustaceans, this autotomy plane is characterized by the presence of a connective tissue sheath and autotomy diaphragm the function of which is to minimize the loss of body fluids when a limb is autotomized (Skinner and Cook, 1991). The observed growth of the blastema, starting within the distal most remaining limb segment and then extending beyond this within a protective cuticle is observed in the Chelicerata, Limulus, and the Crustacea (Gross, 1969; Maruzzo, et al., 2005; Maruzzo and Bortolin, 2013). The presence of regenerating limbs with all segments present but smaller than the corresponding limbs on the same animal as seen in this study, indicates that regeneration requires more than one molt cycle to produce a fully formed and normal size regenerated walking limb (Gross, 1969; Maruzzo, et al., 2005; Maruzzo and Bortolin, 2013). The total number of specimens of Pentapycnon charcoti was relatively small, 10 animals. There was an average of 40% of the specimens from the other three species displaying some degree of regeneration. We could therefore expect about four animals showing some degree of regeneration. The absence of any indication of regeneration including blastema formation may be simply due to the small sample size. Alternately it may be that this genus lacks the ability to autotomize and regenerate limbs. Consequently those individuals losing a limb or part of a limb simply do not survive the trauma. Adult animals which have reached their final size and have stopped molting do not show evidence of regenerative ability according to Fage (1949) and Helfer and Schlottke (1935). In this study evidence of regeneration in adult N. australe in the form of blastema formation was observed. Our results agree with Fleming et al. (2007) in that only limb regeneration occurs in the arthropods. The ability to autotomize damaged limbs and regenerate a replacement has obvious selective advantages for any animal (Maginnis, 2006). Regeneration has been in the ovigerous appendages of C. tortipalpus and the cheliphores, N. australe respectively. Chelophore regeneration had been inferred previously by Child (1979). Acknowledgements Austin Patrick Harlow is acknowledged for assistance with the photomicrographs. References Bain BA. 2003. Larval types and a summary of post-embryonic development within the Pycnogonida. Invertebrate Reproduction and Development, 43(3): 193-222 Bely AE, Nyberg KG. 2009. Evolution of animal regeneration: re-emergence of a field. Trends in Ecology and Evolution, 25(3): 161-170 Child CA. 1979. Shallow-Water Pycnogonida of the Isthmus of Panama and the Coasts of Middle America. Smithsonian Institution Press, Washington DC, USA Dohrn A. 1881. Die Pantopoden des Golfes von Naples. Fauna und Flora des Golfo von Naples, Monograph 3 Fage L. 1949. Classe de Pycnogonides. Traite de Zoologie, 6: 906-941 IAEES www.iaees.org Arthropods, 2019, 9(4): 110-117 117 Fleming PA, Muller D, Bateman PW. 2007. Leave it all behind: a taxonomic perspective of autotomy in invertebrates. Biological Reviews, 82: 481-510 Gaubert P. 1881. Automie chez les Pycnogonides. Bulletin Society Zoological France, 17: 224-225 Helfer H, Schlottke E. 1935. Pantopoda. Dr. H. G. Bronns Kl OrdnTierreichs 5: 1-314 King PE. 1973. Pycnogonids. St Martin’s Press. New York, USA Krishnakumaran A, Schniderman HA. 1970. Control of molting in mandibulate and chelicerate arthropods by ecdysones. Biological Bulletin, 139(3): 420-538 Loeb J. 1905. Studies in general Physiology. The University of Chicago Press, USA Maginnis TL. 2006. The costs of autotomy and regeneration in animals: a review and framework for future research. Behavioral Ecology, 17: 857-872 Morgan TH. 1901. Regeneration and liability to injury. Science, 14(346): 235-248 Maruzzo D, Bonato L, Brena C, Fusco G, Minelli A. 2005. Appendage loss and regeneration in arthropods: A comparative view. In: Crustacea and Arthropod Relationships (Koenemann S, Jenner RA, eds). CRC Press, USA Maruzzo D, Bortolin F. 2013. Arthropod regeneration. In: Arthropod Biology and Evolution. (Minelli A, Boxshall G, Fusco G, eds). Springer, London, UK Mitić BM, Tomić VT, Makarov SE, Ilić BS, Ćurćić BPM. 2010. On the appendage regeneration of Eupolybothrus transsylvanicus (Latzel) (Chilopoda: Lithobiidae). Archives of Biological Science, 53: 2122 Needham AE. 1952. Regeneration and Wound Healing. John Wiley & Sons, New York, USA Skinner DM, Cook J. 1991. New limbs for old: some highlights in the history of regeneration in crustacea. In: A History of Regeneration Research: Milestones in the Evolution of a Science (Densmore CE, ed). 25-45, Cambridge University Press, New York, USA IAEES www.iaees.org Arthropods, 2019, 8(4): 118-126 Article First record of the crab, Droippe quadridens (Fabricius, 1793) (Brachyura: Dorippidae), from the Iraqi coastal waters of the NW Arabian Gulf, with notes on the occurrence of seven species of crabs in the region KhaledKh Al-Khafaji1, Tariq H.Y. Al-Maliky1, Anwar M.J. Al-Maliky2 1 Marine Biology Department, Marine Science Centre, Basrah University, Iraq 2 Biology Department, College of Education, Basrah University, Iraq Email: Khaledalkhafaji70@gmail.com Received 20 August 2019; Accepted 25 September 2019; Published 1 December 2019 Abstract The present study is important in concerning the ecological and classification of the native and invasive species to the Iraqi coast in the north-west of the Arabian Gulf. It has been noted that there is a recent trend to record many decapod crustaceans species in general and marine crabs in particular. The aim of the presence article is to find out diversity and distribution of the dorippid crabs (family: Dorippidae) in addition to other brachyuran crabs species in the subtidal zone along the Iraqi coast. The present study was conducted in summer and winter months from April 2016 to March 2017. Three sites for collection of samples along the Iraqi coast were selected, site 1 in Khor Abudallah canal, site 2 in the coastal region around south Al-Fao town, and site 3 in Rass Al-Besha area. Sampling was carried out using a shrimp’s trawler nets. One of the important results of this study is that specimens of the crab Dorippe quadridens (Fabricius, 1793) were recorded for the first time along the Iraqi coasts. During this survey, seven species of other Brachyuran crabs were identified; belonging to the seven families. The study recommended continuous monitoring of the brachyuran crabs and other invertebrates species that inhabiting the Iraqi coast in order to provide basic information on the species diversity and distribution of marine crabs inhabiting this harsh environment. Keywords Khor Abudallah; Rass Al-Besha; crab; Dorippe quadridens; diversity; distribution. Arthropods ISSN 2224­4255 URL: http://www.iaees.org/publications/journals/arthropods/online­version.asp RSS: http://www.iaees.org/publications/journals/arthropods/rss.xml E­mail: arthropods@iaees.org Editor­in­Chief: WenJun Zhang Publisher: International Academy of Ecology and Environmental Sciences 1 Introduction Most of research about the taxonomic diversity of decapod crustaceans have increased considerably during the last decades (Ng et al., 2008; Clores and Ramos, 2013; Varadharajan et al., 2013; Al-Maliky et al., 2016; AlKhafaji et al., 2017). Many of these researches focused on decapod crustaceans inhabiting marine ecosystems, IAEES www.iaees.org Arthropods, 2019, 8(4): 118-126 119 whereas the diversity, distribution and abundance of marine crab species are one among the issues (Zairion et al., 2018). Several previous articles on D. quadridens which is belonging to the family Dorippidae were focused on the occurrence and distribution of this species and other dorippid crabs in their habitats including the coasts of Australia (Davie, 2002); coastal waters of Madagascar, New Caledonia, Indonesia and the Philippines (Chen, 1987, 1993); coastal waters of Cambodia (Jensen et al., 2011). The main cause of change in biodiversity may be the occurrence of climate change phenomena, such as high temperatures and low rainfall, and thus migration of these invasive species from their original environments to our environments (Zhang and Chen, 2011). The specimens of this species were found and recorded in previous studies in the west, east and south coasts of the Arabian Gulf. However, there are limited information on the dorippid crabs and other brachyuran crabs in the Arabian Gulf waters, including their diversity, distribution and notes on the occurrence of these species in Arabian Gulf waters (Stephensen, 1946; Naderloo and Sari, 2007; Naderloo and Turkey, 2012; Naderloo et al., 2015). The aim of the present study is to increase the knowledge on the marine fauna of the Iraqi coasts, northwest the Arabian Gulf, which remains incomplete. To increase our understanding of the fauna of this area and to determine the diversity and distribution of the dorippid crabs: D. quadridens (Fabricius, 1793) in the subtidal zone along the Iraqi coast, NW Arabian Gulf, Iraq. 2 Materials and Methods Specimens of the dorippid species and some other Brachyuran crabs species from the subtidal zone were collected from three main sites during the summer and winter months at the period from April 2016 to March 2017. Three stations were selected and sampled once by shrimps trawl net, each trawl lasted for a maximum of 30 min. Three stations in Iraqi coastal waters at south of Al-Basrah city, were sampled: (1) site 1, Khor Abdullah coast; (2) site 2, in coastal waters around the south of Al-Fao town, and (3) site 3, from the shallow subtidal zone at Rass Al-Bessha coastal region at the southern end of the Shatt Al-Arab at Al-Fao town (Fig. 1). The depth at these stations was ranging from 5-25 m. Samples of the crabs were brought to the laboratory at the Marine Biology Dep. of the Marine Science Center. The species were identified with the aid of the following: Stephensen (1946), Naderloo and Sari (2007), and Naderloo and Turkey (2012). Specimens of each species were placed in a plastic container with label and preserved by 70-80% alcohol. The “winter” season included December through February while the “summer” season included May through July 2016. 3 Results and Discussion Systematic Account Family Dorippidae MacLeay, 1838 Subfamily Dorippinae MacLeay, 1838 Genus Dorippe Weber, 1795 Species Dorippe quadridens (Fabricius, 1793) 3.1 First record of Dorippe quadridens (Fabricius, 1793) (Crustacea, Decapoda, Dorippidae) Dorippoides nudipes (Manning and Holthuis, 1986) is a common species in the subtidal zones of the Arabian Gulf. It is found in variety of habitats including sandy, muddy and rocky substrate, but it was not collected during the present study. The present paper deals with as new records for the Iraqi coast from the subtidal IAEES www.iaees.org 120 Arthropods, 2019, 8(4): 118-126 zones, along the Iraqi coasts, NW-Arabian Gulf. Moreover, it is the first record of D. quadridens (Fabricius, 1793) in the Arabian Gulf. Fig. 1 Sampling sites along the Iraqi coasts. The specimens of D. quadridens (Fabricius, 1793) were collected by bottom trawl net in the Iraqi coastal of water, NW Arabian Gulf in winter 2016 and summer 2017, from the three sites at depths of 5 - 25 m. The specimen of D. quadridens (Fig. 1, a) fits the description of Holthuis and Manning (1990) as: carapace strongly sculptured, granulated, bearing pubescence and grooves evident. Tubercles usually well indicated, sometimes low. Surface covered by long, flexible hairs, worn off in old individuals. Two lateral tubercles on cardiac area. Antero-lateral margin of carapace, between base of exorbital tooth and cervical groove, with few to many (3-9) sharp denticles. Front teeth flat, with narrowly rounded apices, separated by a deep but open V. Lower orbital margin with row of 4-7 spines but without additional row of denticles. Carpus of cheliped with distinct spinules and hairs on upper surface; palm of chela smooth, except for granules in extreme proximal part. P2-P3: merus compressed, distinctly higher than wide, covered with pubescence; merus of P3 less than six times as long as high. Measurements: carapace length 3.2 cm; carapace width 2.9 cm; length of cheliped 3.4 cm; body depth1.7 cm. Colour: carapace brownish-grey. Fingers of chelas yellowish brown. Weight: 18 g. 3.2 Habitat D. quadridens is found in the sublittoral, shallow water, substrate shelly sand or mud to sand bottoms (Holthuis and Manning, 1990; Thoma, 2007). Dorippid crabs are commonly live in sandy, muddy and rocky substrate at shallow coastal water. D. quadridens was caught in depths of 5 to 40 m (Thoma, 2007), varies from 1 to 73 m and most commonly occurs in 1 to 30 m and ever found (1 individual) in 415 m (Holthuis and Manning, 1990). In this study, we found D. quadridens at a depth of 5 to 25 m. 3.3 Distribution D. quadridens has a wide distribution within the Indo-West Pacific region, extending from the Arabian Gulf to the Red Sea, Suez Canal and eastern Africa. IAEES www.iaees.org Arthropods, 2019, 8(4): 118-126 121 D. quadridens (Fabricius, 1793) is known from Iran (Stephensen 1946, Naderloo and Sari 2007), Kuwait and UAE (Apel, 2001), Saudi Arabia [Basson et al, (1977) as Dorippe dorsipes, Apel (2001)], Bahrain (Stephensen, 1946), Gulf of Oman (Naderloo et al, 2015), Red sea, Pakistan, southern India, southern China, Vietnam, Thailand, Malaysia, Indonesia (Holthuis and Manning, 1990), Cambodia, Madagascar (Chen, 1987), Australia (Manning, 1993; Thoma, 2007), Thailand (Ng and Davie, 2002), New Caledonia, Philippines (Chen, 1993) and now in the subtidal zone of the NW Arabian Gulf Iraqi coasts. The doripped crabs are a macro-benthos, mostly found in tropical waters, and have no economic value and consumption. Familly Dorippidae consists of nine genera, two of which are Dorippe and Dorippoides. There are seven species of the genus Dorippe (Manning, 1993) and 2 species of the genus Dorippoides (Holthuis and Manning, 1990) are known from the Indo-West Pacific region. Specimens from other species of brachyuran crabs were collected and identified during the present survey (Fig. 2) (Table 1), shows a list of the recorded species at the three stations. Other brachyuran were represented by seven genera, seven families Epialtidae, Leucosiidae, Galenidae, Portunidae, Xanthidae, Varunidae and Matutidae, each comprised of only one species. Table 1 List of the other brachyuran crab species recorded in the present surveywith details of records by earlier workers. Family 1-Xanthidae 2-Matutidae 3- Varunidae 4- Portunidae Species list Atergatis roseus (Rüppell, 1830) Matuta planipes Fabricius, 1798 Metaplax indica (H. Milne Edwards, 1852) Portunus pelagicus (Linnaeus, 1758) Recordby Al-Khafaji et al. (2017) Al-Khafaji et al. (2017) Al Maliky et al. (2016) Al-Khafaji et al. (2017) 5-Leucosiidae Hiplyra sagitta (Galil, 2009) Al-Khafaji et al. (2017) 6-Galenidae Halimedetyche (Herbst, 1801) Al-Khafaji et al. (2017) 7-Epialtidae Hyastenus hilgendorfi (De Man, 1887) Al-Khafaji et al. (2017) Table 2 The presence and absence of the species recorded at each site during the survey in the present study (+: present, -: absent, S: summer, W: winter). No Family Species list Site 1 1 1-Xanthidae Atergatis roseus S + W S + - W - S - W - 2 2-Matutidae Matuta planipes + + - + - + 3 4 3-Dorippidae 4- Varunidae Dorippe quadridens(Fabricius, 1793) Metaplax indica - - + + + + + + + 5 5- Portunidae Portunus pelagicus + + + + + + 6 6-Leucosiidae Hiplyra sagitta - + - + - + 7 7-Galenidae Halimedetyche + + + + + - 8 Total 8-Epialtidae 8 Hyastenus hilgendorfi 8 - + + + + + IAEES Site 2 Site 3 www.iaees.org 122 Arthropods, 2019, 8(4): 118-126 The presence and absence of the species in winter and summer were recorded at each site during the survey period (Table 2). The blue swimming crab, Portunus pelagicus (Linnaeus, 1758) was the most commonly distributed species caught by shrimp’s trawlers in the subtidal zone from the three sites at the two seasons along the Iraqi coastal waters, NW Arabian Gulf. The spider crabs Hyastenus hilgendorfi De Man, 1887 and Halimedetyche (Herbst, 1801) were the second common species. Hyastenus hilgendorfi was absent only in site 1 in summer, while Halimedetyche was absent only in site 3, Atergatis roseus (Rüppell, 1830) was the least present species caught and recorded only in site 1. a. Dorippe quadridens (Fabricius, 1793) b. Atergatis roseus (Rüppell, 1830) c. Hyastenus hilgendorfi (De Man, 1887) IAEES www.iaees.org Arthropods, 2019, 8(4): 118-126 123 d. Hiplyra sagitta (Galil, 2009) e. Halimedetyche (Herbst, 1801) f. Metaplax indica (H. Milne Edwards, 1852) IAEES www.iaees.org 124 Arthroopods, 2019, 8(4 4): 118-126 g. Matuuta planipes (F Fabricius, 1798) h. Portunuss pelagicus (L Linnaeus, 17588) Fig. 2 a, b, b c, d, e, f, g annd h: (Left imagge -dorsal view w, right image - ventral view). Crab species coollected from th he Iraqi coastall waters of the t NW Arabiaan Gulf. 4 Conclu usion The findding of D. quuadridens (Faabricius, 17933) where the species is widely w distribuuted in the eaast, west andd south of Arabian Gulff, is not surprrising in the subtidal s zone along of Iraqqi coastal waaters, NW of Arabian A gulf,, c easily disstributed by a migration to t another areeas. One morre reason, thee climate andd because this species can mental conditions in the Arabian A Gulf waters is sim milar to that inn Iraqi coastaal waters. In addition, thee environm present study s adds to the brachyuran crabs, onee as new species recorded in Iraqi coasttal waters. wledgments Acknow We wouuld like to thaank the mem mbers of the Marine Biology Departm ment, Marine Sciences Ceenter, Basrahh Universitty, for help me m in accomplishing this work w and for their t valuablee advice and ssuggestions. Referencces Al-Khafaaji KK, Al-W Waeli AA, Al--Maliky TH. 2017. New records r of xaanthid crabs A Atergatis roseeus (Rüppell,, 18300) (Crustacea: Decapoda: Brachyura) B frrom Iraqi coaast, south of Basrah B city, Irraq. Arthropo ods, 6(2): 54-58 IAEES www.iaees.org w Arthropods, 2019, 8(4): 118-126 125 Al-Maliky TH, Naser MD, Yasser AGh, et al. 2016. New record of the Grapsoid crab Metaplax indica H. Milne-Edwards, 1852 (Decapoda: Brachyura: Thoracotremata) from the NW of the Arabian Gulf, Iraq. Arthropods, 5(1): 23-27 Apel M. 2001. Taxonomie und zoogeographie der brachyura, paguridea und porcellanidae (Crustacea: Decapoda) des Persisch-Arabischen Golfes, Dissertation zur Erlangung des Doktorgrades der Naturwissenschaften. Johann Wolfgang Goethe University, Frankfurt am Main, Germany Basson PW, Burchard JE, Hardy JT, Price ARG. 1977. Biotopes of the Western Arabian Gulf: Marine life and environments of Saudi Arabia, Dhahran: ARAMCO, Dept. of Loss Prevention and Environmental Affairs, Saudi Arabia Chen HL. 1987. Dorippidae (Crustacea Decapoda Brachyura) collected in Madagascar waters. Bulletin du Muséum national d'histoire naturelle Paris, 4e Ser., 9, sect. A(3): 677-693 (figs 1-7, pis 1-2) Chen HL. 1993. Crustacea Decapoda: Dorippidae of New Caledonia, Indonesia and the Philippines. Resultats des Campagnes MUSORSTOM (Vol 10) (Crosnier A, ed) (Paris: MemMusnatnHistnat). 315-345, Paris, France Clores M, Ramos GB. 2013. Reproductive characteristics of a brachyuran crab, Grapsus tenuicrustatus (Herbst, 1783) (Decapoda: Grapsidae) found in Talim Bay, Batangas, Philippines. Arthropods, 2(3): 111125 Davie PJF. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda - Anomura, Brachyura. In: Zoological Catalogue of Australia (Wells A, Houston WWK, eds). CSIRO Publishing, 19.3B Melbourne, Australia Holthuis LB, Manning RB. 1990. Researches on Crustacea (Special Number 3). The Carcinological Society of Japan, Tokyo, Japan Jensen KR, Ing T, Va L. 2011. First record of the rare porcellanid crab Pseudoporcellanella manoliensis Sankarankutt y, 1961 (Crustacea: Anomura) in the coastal waters of Cambodia. Cambodian Journal of Natural History, 2: 81-85 Manning RB. 1993. Two New Dorippid Crabs from Australia (Crustacea: Decapoda: Dorippidae Rec. Australian Museum, 45: 1-4 Manning RB, Holthuis LB. 1981. West African brachyuran crabs (Crustacea: Decapoda). Smithsonian Contributions to Zoology, 306: 1-379 Naderloo R, Sari A. 2007. Subtidal crabs of the Iranian coast of the Persian Gulf: new collections and biogeographic consideration, School of Biology, University College of science, University of Tehran, Iran. Aquatic Ecosystem Health & Management, 10(3): 341-349 Naderloo R, Türkay M. 2012. Decapod crustaceans of littoral and shallow sublittoral habitats along the eastern (Iranian) coast of the Persian Gulf: faunistics, biodiversity and zoogeography. Zootaxa, 3374: 1-67 Naderloo R, Ebrahimnejad S, Sari A. 2015. Annotated checklist of the decapod crustaceans of the Gulf of Oman, northwestern Indian Ocean. Zootaxa, 4028: 397-412 Ng PKL, Davie PJF. 2002. A checklist of the brachyuran crabs of Phuket and western Thailand. Phuket Marine Biological Center Research Bulletin, 23: 369-384 Ng PKL, Guinot D, Davie PJF. 2008. Systema Brachyurorum: Part I. An annotated checklist of extant brachyuran crabs of the world The Raffles. Bulletin of Zoology, 17: 1-286 Stephensen K. 1946. The Brachyura of the Iranian Gulf, Danish Scientific Investigations in Iran, Part IV. E. 57-237, Munksgaard, Copenhagen, Denmark Thoma BT. 2007. Notes on crabs of the families Goneplacidae and Dorippidae (Decapoda: Brachyura) from the Dampier Archipelago, Western Australia. Records - Western Australian Museum, 73: 299-302 IAEES www.iaees.org 126 Arthropods, 2019, 8(4): 118-126 Thomassin BA. 1974. Soft Bottom Carcinological Fauna Sensulata On TulÉAr Coral Reef Complexes (S.W. Madagascar): Distribution, Importance, Roles Played in Trophic Food-Chains and in Bottom Deposits (Vol 1). 297-320, Great Barrier Reef Committee, Brisbane, Australia Varadharajan D, Soundarapandian P, Pushparajan N. 2013. The global science of crab biodiversity from Puducherry coast, south east coast of India. Arthropods, 2013, 2(1): 26-35 Zairion AA, Hakim A, Mashar A, et al. 2018. Diversity and distribution of Dorippid crabs (Brachyura: Dorippidae) in east coast of Lampung, Indonesia. IOP Conference Series: Earth and Environmental Science, 149: 012056 Zhang WJ, Chen B. 2011. Environment patterns and influential factors of biological invasions: a worldwide survey. Proceedings of the International Academy of Ecology and Environmental Sciences, 1(1): 1-14 IAEES www.iaees.org Arthropods, 2019, 8(4): 127-136 Article Xylophagous millipede surface area to volume ratios are sizedependent in forests Mark Cooper School of Animal, Plant & Environmental Sciences, University of the Witwatersrand, Johannesburg 2050, South Africa Email: cm.i@aol.com Received 14 August 2019; Accepted 20 September 2019; Published 1 December 2019 Abstract A consistent effect of increasing precipitation (and resource abundance) on body size reductions is known as a water conservation hypothesis. Here a water conservation hypothesis was investigated in millipedes and a comparison made between high long-term mean annual precipitation of forest (750-1500 mm) and lower longterm mean annual precipitation of savanna (544 mm) biome species (n=29, 6). When the confounding effects of phylogeny, sexual dimorphism, sexual size dimorphism and size were controlled/removed, differences were found between six savanna species (Bicoxidens brincki, Doratogonus annulipes, Harpagophora spirobolina, Julomorpha hilaris, J. panda, Odontopyge tabulinus: 0,35975-2,632336 mm-1) and 29 forest species (Centrobolus: 0,000113-0,679931 mm-1; Sphaerotherium: 1,14271-3 mm-1) in the surface area: volume ratios. Savanna millipedes had size-independent surface area: volume ratios (0,519783 mm-1 in males and 0,823878 mm-1 in females). Differences occurred between size-independent savanna and size-dependent forest taxa in surface area: volume ratios (t=3.75191, p=0.000013, n=58,12) controlling for the derivation whereby length/width increase affected surface area equally. Female savanna millipedes were longer than female forest millipedes (t=2.26165, p=0.016156, n=22, 6). Keywords area; Centrobolus; conservation; Sphaerotherium; surface; volumes. Arthropods ISSN 2224­4255 URL: http://www.iaees.org/publications/journals/arthropods/online­version.asp RSS: http://www.iaees.org/publications/journals/arthropods/rss.xml E­mail: arthropods@iaees.org Editor­in­Chief: WenJun Zhang Publisher: International Academy of Ecology and Environmental Sciences 1 Introduction There was a consistent effect of increasing precipitation (and resource abundance) on body size reductions of an entire order of legless, predominantly underground‐dwelling amphibians (Gymnophiona, or caecilians), supporting the water conservation hypothesis (Lees, 1950; Pincheira-Donoso et al., 2019). The humidity (‘water conservation hypothesis’) “rests on three conditions: that spiracular transpiration is greater than cuticular transpiration; that cuticular transpiration rates are lower in desert species; and that changes in body form associated with flightlessness lead to an overall reduction in water loss rates. The extreme form of the IAEES www.iaees.org 128 Arthropods, 2019, 8(4): 127-136 morphological-convergence condition suggests that this change in body shape should be most pronounced in desert-dwelling taxa” such as beetles (Chown et al., 2011). Here the water conservation hypothesis was investigated in millipedes. Smaller millipedes, having lower water reserve, higher cuticular permeability values and a higher rate of per cent of total body water loss, were found to be less tolerant to desiccation compared with larger species (Bhakat, 2014). Water relations in the desert millipede Orthoporus ornatus is considerably greater than in millipedes previously studied (Crawford, 1972). The percentage of total body water loss increases linearly with desiccation time in the garden millipede Oxidus gracilis (Appel, 1988). Water is readily lost and taken up through the cuticle, the effect of the spiracles and of excretion being negligible (Cloudsley-Thompson, 1950). It was also noted the percentage water content of smaller millipedes is greater than larger ones (Baker, 1980). Here a comparison was made between millipede species of forest and savanna biomes (Geldenhuys, 1989; Kulmatiski and Beard, 2013). When the confounding effects of sexual size dimorphism were removed, differences were investigated between species of the savanna species and their forest counterparts in surface area to volume ratios. 2 Materials and Methods 2.1 Morphometrics calculations Body volumes, surface areas and surface area to volume ratios were calculated in 28 forest species compared to 6 savanna species. Two morphometric parameters were used to obtain measurements, length and width, both of which were obtained from the published literature (Cooper, 2018; Cooper, 2019; Lawrence, 1967; Schubart, 1966) (Table 1). Body volumes were calculated based on the formula for a cylinder V = πr2h and surface areas were calculated based on the formula for the same cylinder SA = 2πr(r+h) in all species except Sphaerotherium pill millipedes where the body volume formula was V = 4πr3/3 and surface area was SA = 4πr2. 2.2 Statistical tests Body volumes, surface areas and surface area to volume ratios of male and female millipedes for the 28 forest and 6 savanna species were tabulated using a Microsoft Excel spreadsheet. The One-Way ANOVA was performed using summary data to test for differences between taxa using a Free Statistics Calculator version 4.0 available at https://www.danielsoper.com/statcalc/calculator.aspx?id=43. Values were then compared using the http://www.socscistatistics.com website t-test for 2 independent means. Males and females were compared with respect to body volumes, surface areas and surface area to volume ratios across the forest and savanna biomes. Then males were added to females and forest genera (Table 2) compared to each other and to savanna taxa which were pooled (Table 3). 2.3 Control I controlled for the confounding effects of phylogeny, sexual dimorphism, sexual size dimorphism and size in each comparison. 2.4 Environmental variables The long-term mean annual precipitation in the savanna was recorded at 544 mm (Kulmatiski and Beard, 2013), and in the forest was estimated at 750-1500 mm (Geldenhuys, 1989). 3 Results 3.1 Linear measurements Savanna millipedes differed in length compared to forest millipedes (ANOVA: F=2.897, d.f.=3, P=0.042). Savanna millipedes differed in width compared to forest millipedes (ANOVA: F=16.200, d.f.=3, P=0.000). IAEES www.iaees.org Arthropods, 2019, 8(4): 127-136 129 Female savanna helminthomorph millipedes differed from female forest helminthomorph millipedes in length (t=-2.35263, p=0.013091, n=22, 6) but not width (t=1.67428, p=0.053032, n=22, 6). Male savanna helminthomorph millipedes did not differ from male forest helminthomorph millipedes in length (t=-1.40834, p=0.085673, n=22, 6) or width (t=0.79823, p=0.215983, n=22, 6). Female savanna millipedes did not differ from female forest millipedes in width (t=1.61841, p=0.057547, n= 29, 6). Female savanna millipedes were different from female forest millipedes in length (t=-2.26165, p=0.016156, n=22, 6). 3.2 Volumes Forest taxa were indifferent to savanna taxa in volume (t=-1.18061, p=0.120627, n=58, 12; ANOVA: F=2.586, d.f.=3, P=0.060). Centrobolus males differed from females in volume (t=2.19256, p=0.016965, n=22, 22). Sphaerotherium males were marginally different from females in volume (t=-1.76762, p=0.05126, n= 7, 7). Centrobolus males differed from Sphaerotherium males in volume (t=4.15584, p=0.000146, n=22, 7). Centrobolus females did not differ from Sphaerotherium females in volume (t=2.52508, p=0.008874, n=22, 7). A combination of the forest taxa i. e. Centrobolus and Sphaerotherium differed between sexes (ANOVA: F=5.081, d.f.=1, P=0.028). Forest males were no different to savanna males (t=0.26026, p=0.398016, n=35, 6) while forest females were not different to savanna females (t=-1.28835, p=0.102608, n=35, 6). 3.3 Surface areas A difference was present among forest and savanna millipede surface areas (ANOVA: F=341,864.807, d.f.=3, P=0.000). Centrobolus males did not differ from females in surface area (t=-1.24616, p=0.108044, n=22, 22). Sphaerotherium males were marginally different from females in surface area (t=-1.75744, p=0.5215, n=7, 7). Forest taxa did not differ from savanna taxa in surface area (t=-0.32209, p=0.374262, n=58, 12). When sexual size dimorphism was controlled (and Sphaerotherium excluded) in a comparison between Centrobolus and savanna taxa no difference was found in the surface areas (t=0.45811, p=0.32389, n=44, 12). 3.4 Surface area to volume ratios When the forest data set was compared with the savanna (males and females added) a significant difference was found in surface area to volume ratios (t=-3.75191, p= 0.000013, n=58, 12; ANOVA: F=12,927,853.340, d.f.=3, p=0.000). Male and female Centrobolus surface area to volume ratios were not significantly different (t=0.44722, p=0.327921, p<0.10, n=22, 22). The same was true for Sphaerotherium male and female surface area to volume ratios which were indifferent (t=-0.32315, p=0.749501, p<0.10, n=7, 7). Because of this, differences were investigated between species belonging to forest and savanna biomes and there were no differences between male savanna millipede surface area to volume ratios and male forest millipede surface area to volume ratios (t=-2.44161, p=0.008655, n=28, 6). There was a significant difference between forest millipede female surface area to volume ratios and savanna millipede female surface area to volume ratio (t=2.83273, p=0.003045, n=28, 6) but there was no difference between male and female savanna millipede surface area to volume ratios (t=-0.47794, p=0.637407, n=6, 6). When I controlled for phylogeny differences occurred between the surface area to volume ratios of female forest Centrobolus and female (t=-3.39958, p=0.000638, n=22, 6) and male (t=-2.43107, p=0.009202, n=22, 6) savanna millipedes but not with the Sphaerotherium males (t=-1.77194, p=0.041023, n=22, 6) or females (t=-2.23741, p=0.014665, n=22, 7). Male forest Centrobolus differed from female savanna millipedes (t=3.96105, p=0.00011, n=22, 6) and male savanna millipedes (t=-2.88846, p=0.00278, n=22, 6) in surface areas. When sexual size dimorphism was controlled and the sexes were added, there was a difference between the surface area to volume ratios of Centrobolus and Sphaerotherium (t=-3.22188, p=0.000836, n=44, 14). IAEES www.iaees.org Arthropods, 2019, 8(4): 127-136 130 . Species Table 1 Male and female morphometric parameters recorded in savanna and forest millipedes. Male length (mm) Male width (mm) Female length (mm) Female width (mm) B. brincki 93 5.9 84 5.9 C. albitarsus 39 4.0 50 6.0 C. anulatus 69 5.3 76 5.9 C. decoratus 43 4.5 31 4.2 C. digrammus 41 4.0 34 4.4 C. dubius 52 5.0 51 5.9 C. fulgidus 54 5.2 52 6.8 C. immaculatus 49 4.7 60 7.0 C. inscriptus 67 5.9 63 6.7 C. inyanganus 40 4.5 43 5.2 C. lawrencei 43 4.7 43 5.9 C. lugubris 53 6.2 63 8.4 C. promontorius 33 3.6 27 3.3 C. pusillus 39 4.0 40 5.7 C. richardi 59 5.2 50 5.5 C. ruber 58 5.0 62 6.1 C. rugulosus 49 5.4 50 7.5 C. sagatinus 49 6.2 48 7.0 C. silvanus 46 4.4 44 4.8 C. titanophilus 28 4.1 29 4.3 C. transvaalicus 39 4.4 38 5.0 C. tricolor 45 4.5 37 5.2 C. vastus 65 6.0 63 8.2 D. annulipes 104 5.5 89 5.9 H. spirobolina 72 4.4 79 5.9 J. hilaris 26 2.4 28 3.7 J. panda 32 4 38 2.7 O. tabulinus 63 5 70 5 S.cinctellum 15.5 18.6 S. commune 6 9.5 S. punctulatum 12 21 S. spinatum 11.5 15 S. tenuitarse 7 8 S. trichopygum 10.75 16.5 S. tuberosum 6.75 9 IAEES www.iaees.org Arthropods, 2019, 8(4): 127-136 131 Table 2 Surface area to volume ratios for forest millipedes (Centrobolus, Sphaerotherium). Species Male volume Male surface Female 3 2 volume (mm ) area (mm ) 3 Female Male surface Female surface surface area area: volume area: volume 2 (mm ) -1 (mm ) (mm ) (mm-1) C. albitarsus 1960 5655 1 080,708 2 111,15 0,00051 0,000177 C. annulatus 2058 1729 2462,874 3026,009 0,000486 0,000578 C. decoratus 2736 1718 1 343,031 928,906 0,000365 0,54069 C. digrammus 2061 2068 1 130,973 1 061,607 0,000485 0,000484 C. dubius 4084 5577 1 790,708 2 109,328 0,000245 0,000179 C. fulgidus 4587 7554 1 934,216 2 512,269 0,000218 0,000132 C. immaculatus 3400 9236 1 585,813 2 946,814 0,000294 0,000108 C. inscriptus 7327 8885 2 717,289 2 934,185 0,000136 0,000113 C. inyanganus 2545 3653 1 258,208 1 574,818 0,000393 0,000274 C. lawrencei 2984 4702 1 408,627 1 812,762 0,000335 0,000213 C. lugubris 6400 13965 2 306,18 3 768,403 0,000156 0,000716 C. promontories 1343 924 827,872 628,256 0,616435 0,679931 C. pusillus 1960 4083 1 080,708 1 636,707 0,00051 0,000245 C. richardi 5012 4752 2 098,579 1 917,942 0,418711 0,403607 C. ruber 4555 7248 1 972,92 2 621,596 0,00022 0,000138 C. rugulosus 4489 8836 1 845,749 2 709,624 0,000223 0,000113 C. sagatinus 5913 7389 2 150,357 2 419,026 0,000169 0,000135 C. silvanus 2798 3185 1 393,359 1 471,773 0,000357 0,000314 C. titanophilus 1479 1685 826,93 899,689 0,559114 0,53394 C. transvaalicus 2372 2985 1 199,837 1 350,885 0,000422 0,000335 C. tricolor 2863 3143 1 399,58 1 378,782 0,000349 0,000318 C. vastus 7351 13308 2 676,637 3 668,375 0,000136 0,000751 S. cinctellum 1950 3369 3 019,071 4 347,462 1,548205 1,290294 S. commune 113 449 452,389 1 134,115 4 2,525612 S. punctulatum 905 4849 1 809,557 5 541,769 2 1, 14271 S. spinatum 796 1767 1 661,903 2 827,433 2,08794 1,599887 S. tenuitarse 180 268 615,752 804,248 3,422222 3 S. trichopygum 650 2352 1 452,201 3 421,194 2,233846 1,454507 S. tuberosum 161 381 530,929 1 017,876 3,298137 2,671916 IAEES www.iaees.org Arthropods, 2019, 8(4): 127-136 132 Fig. 1 Size-dependent relationships between the surface area to volume ratios and volume in female forest millipedes (topleft), male forest millipedes (top-right), female Sphaerotherium (lower-left) and male Sphaerotherium (lower-right). All X-values are volume and Y-values are surface area to volume ratios. Surface area to volume ratio was negatively correlated with volume in female Centrobolus (r=0.4577, r =0.2095, n=22, p=0.032499), female Sphaerotherium (r=0.9118, r2=0.8314, n=7, p=0.004324), and female forest millipedes in general (r=-0.5542, r2=0.3071, n=29, p=0.00182). This correlation was not found in male Centrobolus (r=-0.2836, r2=0.0804, n=22, p=0.20189), but was found in male Sphaerotherium (r=-0.8728, r2=0.7618, n=7, p=0.010496) and was found in forest male millipedes in general (r=-0.6529, r2=0.4263, n=29, p=0.000123). Surface area to volume ratios did not correlate with volume in savanna males (r=-0.1538, r2=0.0237, n=6, p=0.770826) or females (r=-0.2131, r2=0.0454, n=6, p=0.685332). 2 IAEES www.iaees.org Arthropods, 2019, 8(4): 127-136 133 Table 3 Surface area to volume ratios of savanna millipedes (Bicoxidens, Doratogonus, Harpagophora, Julomorpha, Orthoporoides). Species Body volume (mm3) Surface area (mm2) Surface area: Volume (mm-1) Male Female Male Female Male Female B. brincki 10 300,656 9 275,598 3 695,762 3 345,919 0,358789 0,360723 D. annulipes 9 899,293 9 858,974 3 789,809 3 546,761 0,382836 0,35975 H. spirobolina 4 391,946 8 639,348 2 111,37 3 147,31 0,480737 0,364299 J. hilaris 474,104 1 195,634 431,278 3 147,31 0,90967 2,632336 J. panda 1 608,495 865,704 904,779 687,066 0,5625 0,79365 O. tabulinus 5 170,96 5 407,625 2 193,341 2 338,853 0,424165 0,43251 4 Discussion Millipede species-specific volumes are known to exist and these correlate with bimaturism, copulation duration, fecundity, female body width, sexual conflict, sexual size dimorphism, species and mass (Cooper, 2016-2019). Forest millipede surface area to volume ratio was size-dependent (Fig. 1) while in savanna millipedes it was size-independent. The significant difference between forest millipede female surface area to volume ratios and savanna millipede female surface area to volume ratio was a finding which suggests differences in the form in agreement with the water conservation hypothesis (Lees, 1950). It suggests there are precipitation-size patterns in worm-like millipedes which may affect the adaptability to and validity of biological rules (Meiri and Dayan, 2003; Schmidt-Nielson, 1984). Although sexual dimorphism is not clearly evident in the savanna biome their size is thought to be mostly longer (Cooper, 2019). Where there is the confounding effect of sexual dimorphism in the forest millipedes there was a relationship between the surface area to volume ratios and volume. When the effect of sexual dimorphism was removed and surface area to volume ratios were compared, savanna and forest taxa showed a significantly different surface area to volume ratio. Surface area to volume ratios was higher in the savanna taxa although forest taxa were also high due to Sphaerotherium. How do millipedes maximize their size – through an increase in width or length of their cylindrical bodies? It was achieved through a change in width and length, which is probably the most powerful way to maximize the volume and surface area to volume ratio of a cylinder, which is anamorphosis (Enghoff, 1993). A further difference within forest millipede surface area to volume ratios between the genera Centrobolus and Sphaerotherium indicate divisions within the forest taxa suggesting there is water conservation stress within the forest as well. The forest genus Sphaerotherium illustrated the most differences with volume, surface area and surface area to volume ratios all being different and a strong relationship between volumes and surface area to volume ratios. This genus is related to Glomeris and the water relations were attributed to size and conglobation (Edney, 1951). Conglobation in the pill bug (Armadillidium vulgare) is an adaptive water conservation mechanism (Smigel and Gibbs, 2008). When phylogeny, sexual size dimorphism, and sexual dimorphism were controlled a difference was found in the surface area to volume ratios between the IAEES www.iaees.org 134 Arthropods, 2019, 8(4): 127-136 forest and savanna sample but not volumes. This was evident in the presence of a relationship between forest surface area to volume ratios and volume which was absent in the savanna taxa. It proves forest millipedes which cannot conglobate also conserve water adaptively through the surface area to volume ratios dependent on size. Smaller juliform millipedes, having lower water reserve, higher cuticular permeability values and higher rate of percent of total body water lost, are known to be less tolerant to desiccation compared with larger species (Bhakat, 2014). Therefore the size-dependent surface area to volume ratios of forest millipedes is predictably less tolerant to desiccation while the size-independent surface area to volume ratios of savanna millipedes are predictably intolerant to desiccation. Water relations in the desert millipede Orthoporus ornatus is considerably greater than in millipedes previously studied (Crawford, 1972). The percentage of total body water loss increases linearly with desiccation time in the garden millipede Oxidus gracilis (Appel, 1988). Water is readily lost and taken up through the cuticle, the effect of the spiracles and of excretion being negligible (Cloudsley-Thompson, 1950). It was also noted the percentage water content of smaller millipedes is greater than larger ones (Baker, 1980). Energy and water balances vary from tropical to desert biomes and can also change temporally which are behaviourally modified and independent of surface area to volume ratios but dependent on genera (Clousley-Thompson, 1959; Crawford, 1978; Webb and Telford, 1995). In order to conserve water, the terrestrial arthropods have also acquired a relatively impervious integument (Dwarakanath and Job, 1965). The significantly higher surface area to volume ratios of female forest millipedes compared to female savanna millipedes suggests a combined effect of fecundity selection together with water conservation. This is seen and probably thought to be caused due to a difference in lengths of the two, which affects the surface area of the cylindrical body form as powerfully as width which was the case. The savanna millipedes had longer females than the shorter forest female millipedes. This suggests millipede body size can change independent to temperature especially in the size-independent savanna millipedes (Enghoff, 1992; Golovatch and Kime, 2009). Behavioural differentiation and different use of time budget may contribute to the trophic niche separation among coexisting millipede species and in this instance sexes because distance passed per day correlates with body length (Semenyuk and Tiunov, 2019). Surface area to volume ratios is affected more through changes in width than length which is seen in the female differences due to fecundity selection (Darwin, 1874). References Appel AG. 1988. Water relations and desiccation tolerance of migrating garden millipedes (Diplopoda: Paradoxosomatidae). Environmental Entomology, 17(3): 463-466 Baker GH. 1980. The water and temperature relationships of Ommatoiulus moreletii (Diplopoda: Iulidae). Journal of Zoology, 190: 97-108 Bhakat S. 2014. Comparative water relations of some tropical millipedes. Kragujevac Journal of Science, 36: 185-194 Chown SL, Pistorius PA, Scholtz C. 2011. Morphological correlates of flightlessness in southern African Scarabaeinae (Coleoptera: Scarabaeidae): Testing a condition of the water conservation hypothesis. Canadian Journal of Zoology, 76: 1123-1133 Cloudsley-Thompson JL. 1950. The water relations and cuticle of Paradesmus gracilis (Diplopoda, Strongylosomidae). Journal of Cell Science, S3-91: 453-464 Clousley-Thompson JL. 1959. Studies in diurnal rhythms ix the water-relations of some nocturnal tropical arthropods. Entomologia Experimentalis et Applicata, 2(4): 249-256 Cooper MI. 2016. Heavier-shorter-wider females in the millipede Centrobolus inscriptus (Attems). Journal of IAEES www.iaees.org Arthropods, 2019, 8(4): 127-136 135 Entomology and Zoology Studies, 4(2): 509-510 Cooper MI. 2017. Allometry of copulation in worm-like millipedes. Journal of Entomology and Zoology Studies, 5(3): 1720-1722 Cooper MI. 2017. Relative sexual size dimorphism in Centrobolus digrammus (Pocock) compared to 18 congenerics. Journal of Entomology and Zoology Studies, 5(2): 1558-1560 Cooper MI. 2017. Relative sexual size dimorphism in Centrobolus fulgidus (Lawrence) compared to 18 congenerics. Journal of Entomology and Zoology Studies, 5(3): 77-79 Cooper MI. 2017. Relative sexual size dimorphism Centrobolus ruber (Attems) compared to 18 congenerics. Journal of Entomology and Zoology Studies, 5(3): 180-182 Cooper MI. 2018. Allometry for sexual dimorphism in millipedes (Diplopoda). Journal of Entomology and Zoology Studies, 6(1): 91-96 Cooper M. 2018. Centrobolus anulatus reversed sexual size dimorphism. Journal of Entomology and Zoology Studies, 6(4): 1569-1572 Cooper M. 2018. Centrobolus size dimorphism breaks Rensch’s rule. Arthropods, 7(3): 48-52 Cooper M. 2019. Size dimorphism in six juliform millipedes. Arthropods, 8(4): 137-142 Crawford CS. 1978. Seasonal water balance in Orthoporus ornatus, a desert millipede. Ecology, 59(5): 9961004 Crawford CS. 1972. Water relations in a desert millipede Orthoporus ornatus (Girard) (Spriostreptidae). Comparative Biochemistry and Physiology Part A: Physiology, 42(2): 521-535 Darwin, C. 1874. The Descent of Man, and Selection In Relation To Sex. John Murray, London, UK Dwarakanath SK, Job SV. 1965. Studies on transpiration in millepedes I. Spirostreptus asthenes Poc., from a Tropical Jungle Near Madurai. Proceedings of the Indian Academy of Sciences, Section B 61(3): 142-146 David J-F, Geoffroy J-J. 2011. Additional moults into ‘elongatus’ males in laboratory-reared Polydesmus angustus Latzel, 1884 (Diplopoda, Polydesmida, Polydesmidae) – implications for taxonomy. ZooKeys, 156: 41-48 Edney EB. 1951. The evaporation of water from woodlice and the millipede glomeris. Journal of Experimental Biology, 28: 91-115 Enghoff H. 1992. Macaronesian millipedes (Diplopoda) with emphasis on endemic species swarms on Madeira and the Canary Islands. Biological Journal of the Linnean Society, 46: 153-161 Enghoff H, Dohle W, Blower JG. 1993. Anamorphosis in millipedes (Diplopoda)—the present state of knowledge with some developmental and phylogenetic considerations. Zoological Journal of the Linnean Society, 109(2): 103-234 Geldenhuys CJ. 1989. Environmental and biogeographic influences on the distribution and composition of the southern Cape forests (Veld type 4). PhD Thesis, Department of Botany, University of Cape Town, South Africa Golovatch SI, Kime RD. 2009. Millipede (Diplopoda) distributions: A review. Soil Organisms, 81(3): 565597 Kulmatiski A, Beard KH. 2013. Woody plant encroachment facilitated by increased precipitation intensity. Nature Climate Change, 3(9): 1-5 Lawrence RF. 1987. The Centipedes and Millipedes Of Southern Africa: Guide. A. A. Bulkeman, Cape Town, South Africa Lees AD. 1950. Water conservation in terrestrial arthropods. Nature, 166: 809-810. Meiri S, Dayan T. 2003. On the validity of Bergmann's rule. Journal of Biogeography, 30(3): 331-351 Pincheira-Donoso D, Meiri S, Jara M, Olalla-Tárraga MA, Hodgson DJ. 2019. Global patterns of body size IAEES www.iaees.org 136 Arthropods, 2019, 8(4): 127-136 evolution are driven by precipitation in legless amphibians. Ecography, DOI:10.1111/ecog.04644 Schmidt-Nielsen K. 1984. Scaling: Why is Animal Size so Important? Cambridge University Press, New York, USA Schubart O. 1966. Diplopoda III. South African Animal Life, 12: 1-227 Semenyuk II, Tiunov AV. 2019. Foraging behaviour as a mechanism for trophic niche separation in a millipede community of southern Vietnam. European Journal of Soil Biology, 90: 36-43 Smigel JT, Gibbs AG. 2008. Conglobation in the pill bug, Armadillidium vulgare, as a water conservation mechanism. Journal of Insect Science, 8(1): 1-9 Webb PI, Telford SR. 1995. Energy and water balance in the large sub-tropical millipede Alloporus bilobatus (Diplopoda: Spirostreptidae). Journal of Insect Physiology, 41(5): 389-393 IAEES www.iaees.org Arthropods, 2019, 8(4): 137-142 Article Size dimorphism in six juliform millipedes Mark Cooper School of Animal, Plant & Environmental Sciences University of the Witwatersrand, Johannesburg 2050, South Africa Email: cm.i@aol.com Received 25 January 2019; Accepted 6 March 2019; Published 1 December 2019 Abstract Sexual Size Dimorphism (SSD) in the diplopod genera Bicoxidens, Doratogonus, Harpagophora, Julomorpha and Orthoporoides has length, width and rings as the main components of interspecific variation. Interspecific variation in size observed in B. bricki Schubart, 1966, D. annulipes Carl, 1917, H. spirobolina (Karsch, 1881), J. hilaris Attems, 1928, J. panda (Attems, 1928) and O. tabulinus (Attems, 1914) and the data sets were tested for normality. Male lengths differed from female lengths in all except J. hilaris which had different widths. Juliform millipedes appear to have decreased in size over evolutionary time and this study presents an interesting finding showing sexual dimorphism based on length in larger species and sexual dimorphism based on width in the smaller species. The reason for this has to do with the constraints imposed through a cylindrical body form which can be changed more powerfully through reducing width rather than length. Keywords diplopod; horizontal; length; tergite. Arthropods ISSN 2224­4255 URL: http://www.iaees.org/publications/journals/arthropods/online­version.asp RSS: http://www.iaees.org/publications/journals/arthropods/rss.xml E­mail: arthropods@iaees.org Editor­in­Chief: WenJun Zhang Publisher: International Academy of Ecology and Environmental Sciences 1 Introduction Diplopoda are important environmental indicators and under-represented in analyses of invertebrate Sexual Size Dimorphism (SSD) which is the condition where the two sexes of the same species exhibit different characteristics beyond the differences in their sexual organs, although common sexual differences are thought to occur in body mass, length, width and leg dimensions of over half the taxa studied (Telford and Dangerfield, 1990; Hopkin and Read, 1992; Barnett and Telford, 1993; Barnett et al., 1993; Telford and Dangerfield, 1993; Barnett and Telford, 1994; Telford and Dangerfield, 1994; Barnett et al., 1995; Webb and Telford, 1995; Aarde et al., 1996; Barnett and Telford, 1996; Telford and Dangerfield, 1996; Telford and Webb, 1998; Cooper, 2014-2019). Diplopods resemble the majority of invertebrates in SSD is mostly reversed (Cooper, 2018). Larger females are thought to result from fecundity selection (Mauritz, 2011). In the present study, SSD in the superorder Juliformia was investigated in B. bricki Schubart, 1966, D. annulipes Carl, 1917, H. spirobolina (Karsch, 1881), J. hilaris Attems, 1928, J. panda (Attems, 1928) and O. tabulinus (Attems, 1914) and 2 factors determining a response in SSD (length and width) tested. IAEES www.iaees.org 138 Arthropods, 2019, 8(4): 137-142 2 Materials and Methods Two factors were obtained from B. bricki, D. annulipes, H. spirobolina, J. hilaris, J. panda and O. tabulinus: (1) body length (mm) (calibrated in mm); and (2) horizontal tergite width (mm). These basic descriptive figures were statistically tested for normality using a Kolmogorov-Smirnov Test Calculator. The D values of length and width was shown from extracted and published data for B. bricki, D. annulipes, H. spirobolina, J. hilaris, J. panda and O. tabulinus (Schubart, 1966). The factors were compared using a t-test for independent means or Mann-Whitney U-test depending on if the data was parametric or non-paramentric, respectively. 3 Results In 6 tests of male and female widths and lengths the following were found: B. brincki Schubart, 1966 Mean male length was 93.28571 mm (SD=10.111286) and was normal (D=0.28694; p=0.52037; n=7). Mean female length was 83.75 mm (SD=18.468119) and was normal (D=0.20156; p=0.84199; n=6). Mean male width was 5.92857 mm (SD=3.173551) and was normal (D=0.25091; p=0.29003; n=7). Mean female width was 5.9375 mm (SD=2.88603) and was normally distributed (D=0.14811; p=0.82514; n=7). Widths were not different (t=0.00807; p=0.496809; n=13). Lengths were significantly different (t=1.79492; p=0.04797; n=13). D. annulipes Carl, 1917 Tests of male and female widths and lengths all were normal. Mean male length was 104.1667 mm (SD=14.972196) and was normal (D=0.19567; p=0.94162; n=6). Mean female length was 89.28571 mm (SD=26.367368) and was normal (D=0.21382; p=0.84574; n=7). Mean male width was 5.5 mm (SD=3.580249) and was normal (D=0.18377; p=0.74842; n=6). Mean female width was 5.92857 mm (SD=2.37403) and was normally distributed (D=0.16272; p=0.79716; n=7). Male length differed from female length (t=1.46963; p=0.084837; n=13). Male widths and female widths were not different (t=-0.32851; p=0.37686; n=13). H. spirobolina (Karsch, 1881) Tests of male and female widths and lengths all were normal. Mean male length was 71.66667 mm (SD=8.062258) and was normal (D=0.30483; p=0.53458; n=6). Mean female length was 79 mm (SD=9.617692) and was normal (D=0.15725; p=0.99764; n=5). Mean male width was 4.41667 mm (SD=2.712206) and was normal (D=0.25676; p=0.34669; n=6). Mean female width was 5,9 mm (SD=3.17805) and was normally distributed (D=0.24239; p=0.52305; n=5). Widths were not different (t=1.26653; p= 0.109937; n=11). Lengths were significantly different (t=-1.61942; p=0.069905; n=11). J. hilaris Attems, 1928 Tests of male and female widths and lengths all were normal. Mean male length was 26.2 (SD=3.563706) and was normal (D=0.2834; p=0.72834; n=5). Mean female length was 27.8 mm (SD=5.215362) and was normal (D=0.21604; p=0.93275; n=5). Mean Male width was 2.4 mm (SD=1.349897) and was normal (D=0.32261; p=0.19988; n=5). Mean female width was 3.7 mm (SD=2.668749) and was normally distributed (D=0.30891; p=0.24086; n=5). Widths were different (t=-1.37457; p=0.093069; n=10). Lengths were not significantly different (t=-0.56639; p=0.29333; n=10). J. panda (Attems, 1928) Tests of male and female widths and lengths all were normal. Mean male length was 32 mm (SD=3.807887) and was normal (D=0.32147; p=0.57919; n=5). Mean female length was 37.8 mm (SD=2.280351) and was normal (D=0.25963; p=0.8136; n=5). Mean Male width was 4 mm (SD=2.905933) and was normal (D=0.26592; p=0.40748; n=5). Mean female width was 2.7 mm (SD=1.418136) and was normally distributed (D=0.28823; p=0.31349; n=5). Widths were not different (t=1.27136; p=0.109891). Lengths were significantly different (t=-2.922; p=0.009616; n=10). IAEES www.iaees.org Arthropods, 2019, 8(4): 137-142 139 O. tabulinus (Attems, 1914) Tests of male and female widths and lengths all were normal. Mean male length was 63.4 375 mm (SD=16.124515) and was normal (D=0.21459; p=0.39631; n=16). Mean female length was 70.25 mm (SD=13.325381) and was normal (D=0.19271; p=0.39686; n=20). Mean male width was 5.09375 mm (SD=1.731719) and was normal (D=0.18187; p=0.21297; n=16). Mean female width was 4.95 mm (SD=2.489851) and was not normally distributed (D=0.2666; p=0.00529; n=20). Widths were not different (U=588; z=0.58362; p=0.28096). Lengths were significantly different (U=103; z=1.79872; p=0.03593). 4 Discussion The normality of length and width of the two sexes in this species is a finding which successfully confirms the analysis of previous data sets and allows predictive power for the morphological data set in the 6 juliform genera. SSD was found based on length only. The finding does not entirely support studies which shows the size of Juliformia “has two main components: body diameter and number of” rings but includes length as a third component of Juliform size (Enghoff, 1992; Ilić et al., 2017). Other correlates of Juliform size include oxygen consumption, precipitation and temperature (Dwarakanath, 1971; Penteado et al., 1991; Echeverría et al., 2014). Size criteria are useful for determining species of Juliformia diplopods (Cooper, 2014-2019). The present research has illustrated what the minimum sizes of data sets need to be in order to be useful for determining sex of juliforms. Acknowledgements The author is grateful to the University of Stellenbosch for Interlibrary loans. References Aarde RJ van, Ferreira SM, Kritzinger JJ. 1996. Millipede communities in rehabilitating coastal dune forests in northern KwaZulu/Natal, South Africa. Journal of Zoology, 238(4): 703-712 Aarde RJ van, Ferreira SM, Kritzinger JJ. 1996. Successional changes in rehabilitating coastal dune communities in northern KwaZulu/Natal, South Africa. Landscape and Urban Planning, 34(3-4): 277-286 Aarde RJ van, Ferreira SM, Kritzinger JJ, Dyk PJ van, Vogt M, Wassenaar TD. 1996. An evaluation of habitat rehabilitation on coastal dune forests in northern KwaZulu-Natal, South Africa. Restoration Ecology, 4(4): 334-345 Barnett M, Telford SR. 1993. The functional morphology of gonopods as evidence for sperm competition in savannah millipedes. In: Abstracts, 9th International Congress of Myriapodology, Paris, France, 2631.07.1993: 11 Barnett M, Telford SR. 1994. The timing of insemination and its implications for sperm competition in a millipede with prolonged copulation. Animal Behaviour, 48(2): 482-484 Barnett M, Telford SR. 1996. Sperm competition and the evolution of millipede genitalia. In: Acta Myriapodologica. Mémoires du Muséum national d'histoire naturelle (Geoffroy JJ, Mauriès JP, Nguyen DuyJacquemin M, eds), N. S. 169: 331-339 Barnett M, Telford SR, Tibbles BJ. 1995. Female mediation of sperm competition in the millipede Alloporus uncinatus (Diplopoda: Spirostreptidae). Behavioral Ecology and Sociobiology, 36(6): 413-419 Barnett M, Telford SR, Villiers CJ de. 1993. Sperm displacement in a millipede? An investigation into the genital morphology of the southern African spirostreptid millipede Orthoporus pyrhocephalus. Journal of Zoology, 231(3): 511-522 IAEES www.iaees.org 140 Arthropods, 2019, 8(4): 137-142 Cooper MI. 2014. Sex ratios, mating frequencies and relative abundance of sympatric millipedes in the genus Centrobolus Cook. Arthropods, 3(4): 174-176 Cooper MI. 2014. Sexual size dimorphism and corroboration of Rensch’s rule in Chersastus millipedes (Diplopoda: Trigoniulidae). Journal of Entomology and Zoology Studies, 2(6): 264-266 Cooper MI. 2015. Competition affected by re-mating interval in a myriapod. Journal of Entomology and Zoology Studies, 3(4): 77-78 Cooper MI. 2015. Elaborate gonopods in the myriapod genus Chersastus. Journal of Entomology and Zoology Studies, 3(4): 235-238 Cooper M. 2016. Post-insemination associations between males and females in Diplopoda. Journal of Entomology and Zoology Studies, 4(2): 283-285 Cooper MI. 2016. Confirmation of four species of Centrobolus Cook based on gonopod ultrastructure. International Journal of Entomology Research, 1(3): 7-9 Cooper MI. 2016. Confirmation of four species of Centrobolus Cook based on gonopod ultrastructure. Journal of Entomology and Zoology Studies, 4(4): 389-391 Cooper MI. 2016. Do females control the duration of copulation in the aposematic millipede Centrobolus inscriptus? Journal of Entomology and Zoology Studies, 4(6): 623-625 Cooper MI. 2016. Fire millipedes obey the female sooner norm in cross mating Centrobolus Cook. Journal of Entomology and Zoology Studies, 4(1): 173-174 Cooper MI. 2016. Gonopod mechanics in Centrobolus Cook. Journal of Entomology and Zoology Studies, 4(2): 152-154 Cooper MI. 2016. Heavier-shorter-wider females in the millipede Centrobolus inscriptus (Attems). Journal of Entomology and Zoology Studies, 4(2): 509-510 Cooper MI. 2016. Instantaneous insemination in the millipede Centrobolus inscriptus (Attems) determined by artificially terminated mating. Journal of Entomology and Zoology Studies, 4(1): 487-490 Cooper MI. 2016. Sexual bimaturism in the millipede Centrobolus inscriptus (Attems). Journal of Entomology and Zoology Studies, 4(3): 86-87 Cooper MI. 2016. 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Journal of Entomology and Zoology Studies, 6(1): 613-616 Cooper MI. 2018. Sexual size dimorphism and the rejection of Rensch’s rule in Diplopoda (Arthropoda). Journal of Entomology and Zoology Studies, 6(1): 1582-1587 Cooper M. 2018. Trigoniulid size dimorphism breaks Rensch. Journal of Entomology and Zoology Studies, 6(3): 1232-1234 Cooper M. 2018. A review of studies on the fire millipede genus Centrobolus (Diplopoda: Trigoniulidae). Journal of Entomology and Zoology Studies, 6(4): 126-129 Cooper MI. 2018. Volumes of Centrobolus albitarsus (Lawrence, 1967). International Journal of Entomology Research, 3(4): 20-21 Cooper M. 2018. Centrobolus anulatus reversed sexual size dimorphism. Journal of Entomology and Zoology Studies, 6(4): 1569-1572 Cooper M. 2018. Centrobolus size dimorphism breaks Rensch’s rule. Arthropods, 7(3): 48-52 Cooper M. 2018. Centrobolus dubius (Schubart, 1966) monomorphism. International Journal of Research Studies in Zoology, 4(3): 17-21 Cooper M. 2018. Centrobolus lawrencei (Schubart, 1966) monomorphism. Arthropods, 7(4): 82-86 Cooper M. 2018. Centrobolus sagatinus sexual size dimorphism based on differences in horizontal tergite width. Journal of Entomology and Zoology Studies, 6(6): 275-277 Cooper M. 2018. Allometry in Centrobolus. Journal of Entomology and Zoology Studies, 6(6): 284-286 Cooper M. 2018. Centrobolus silvanus dimorphism based on tergite width. Global Journal of Zoology, 3(1): 003005 Cooper M. 2019. A review on studies of behavioural ecology of Centrobolus (Diplopoda, Spirobolida, Pachybolidae) in southern Africa. Arthropods, 8(1): 38-44 Cooper MI, Telford SR. 2000. Copulatory sequences and sexual struggles in millipedes. Journal of Insect IAEES www.iaees.org 142 Arthropods, 2019, 8(4): 137-142 Behavior, 13(2): 217-230 David J-F. 1995. Size criteria for the distinction between Cylindroiulus londinensis (Leach) and Cylindroiulus caeruleocinctus (Wood) (Diplopoda: Julidae). Journal of Natural History, 29(4): 983-991 Dwarakanath SK. 1971. The influence of body size and temperature upon the oxygen consumption in the millipede, Spirostreptus asthenes (Pocock). Comparative Biochemistry and Physiology Part A: Physiology, 38(2): 351-358 Echeverría KS, Ignacio C, Bueno-Villegas J. 2014. Relationship between millipede body size (Polydesmida: Xystodemidae: Rhysodesmus) and altitude, latitude, precipitation and temperature. 16th International Conference of Myriapodology Enghoff H. 1992. The Size of a Millipede. Berichte der naturhistorisch-medizinischen Vereins Innsbruck, Supplement, 10 Hopkin SP, Read HJ. 1992. The Biology of Millipedes. Oxford University Press, Oxford, UK Ilić B, Mitić B, Makarov S. 2017. Sexual dimorphism in Apfelbeckia insculpta (L. Koch, 1867) (Myriapoda: Diplopoda: Callipodida). Archives of Biological Sciences, 69(1): 23-3 Mauritz B. 2011. Ecology of the Namaqua Dwarf Adder, Bitis schneideri. Dissertation. University of the Witwatersrand, South Africa Penteado CHS, Hebling-Beraldo MJA, Mendes EG. 1991. Oxygen consumption related to size and sex in the tropical millipede Pseudonannolene tricolor (Diplopoda, Spirostreptida), Comparative Biochemistry and Physiology Part A: Physiology, 98(2): 265-269 Schubart O. 1966. Diplopoda III. In: South African Animal Life, 12: 1-227 Telford SR, Dangerfield JM. 1990. Manipulation of the sex ratio and duration of copulation in the tropical millipede Alloporus uncinatus: a test of the copulatory guarding hypothesis. Animal Behaviour, 40(5): 984-986 Telford SR, Dangerfield JM. 1990. Sex in millipedes: Laboratory studies on sexual selection. Journal of Biological Education, 24: 233-238 Telford SR, Dangerfield JM. 1993. Mating behaviour and mate choice experiments in some tropical millipedes (Diplopoda: Spirostreptidae). South African Journal of Zoology, 28(3): 155-160 Telford SR, Dangerfield JM. 1993. Mating tactics in the tropical millipede Alloporus uncinatus (Diplopoda: Spirostreptidae). Behaviour, 124(1-2): 45-56 Telford SR, Dangerfield JM. 1993. Millipedes mating systems. In: Abstracts, 9th International Congress of Myriapodology, Paris, France, 26-31.07.1993: 82 Telford SR, Dangerfield JM. 1994. Males control the duration of copulation in the tropical millipede Alloporus uncinatus (Diplopoda: Julida). South African Journal of Zoology, 29(4): 266-268 Telford SR, Dangerfield JM. 1996. Sexual selection in savanna millipedes: products, patterns and processes. In: Acta Myriapodologica - Mémoires du Muséum national d'histoire naturelle (Geoffroy JJ, Mauriès JP, Nguyen DuyJacquemin M, eds), N. S. 169: 565-576 Telford SR, Webb PI. 1998. The energetic cost of copulation in a polygynandrous millipede. Journal of Experimental Biology, 201(11): 1847-1849 Webb PI, Telford SR. 1995. Energy and water balance in the large sub-tropical millipede Alloporus bilobatus (Diplopoda: Spirostreptidae). Journal of Insect Physiology, 41(5): 389-393 IAEES www.iaees.org Arthropods, 2019, 9(4): 143-175 Article Inventory of freshwater arthropods in Pakistan Quddusi B. Kazmi, Farhana S. Ghory Marine Reference Collection & Resource Centre, University of Karachi, Karachi, Pakistan Email: farhanaghory@yahoo.com Received 24 October 2019; Accepted 4 November 2019; Published 1 December 2019 Abstract Inventory of free-living freswater arthropods, with synonymical bibliography, occurring in the Pakistani fresh waters is drawn up, almost entirely from taxonomical literature checked until now. Six hundred and thirty two taxa have been recorded since 1892, of these 239 species and sub-species are of Crustacea, 368 species and sub-species are of Insecta, 25 species and sub-species are of Arachnida as well as few unidentified species. The present inventory is composed of the collated records from the publications of the library and internet. Totally 266 selected publications are arranged as resource references. Keywords inventory; freshwater arthropods; Pakistan. Arthropods ISSN 2224­4255 URL: http://www.iaees.org/publications/journals/arthropods/online­version.asp RSS: http://www.iaees.org/publications/journals/arthropods/rss.xml E­mail: arthropods@iaees.org Editor­in­Chief: WenJun Zhang Publisher: International Academy of Ecology and Environmental Sciences 1 Introduction Our scientific knowledge about freshwater invertebrates, although substantial and useful for many groups, is far less than for the vertebrates. The invertebrates are often used as indicators of the state of streams, rivers, lakes and ponds. Inspite of the facts that the diversity of invertebrate species far outnumbers that of the vertebrate animals particularly the Arthropoda, which contains almost 80% of all animal species, and our fresh water bodies are vast and diverse, our concerns stem from the realization that our knowledge of the diversity and variability of arthropod organisms and the ecosystems in which they occur is woefully incomplete. 2 Materials and Methods Inventory of free-living inland arthropods, with synonymical bibliography, occurring in the Pakistani fresh waters is drawn up, almost entirely from taxonomical literature checked until now. The present inventory is composed of the collated records from the publications of the library and internet. Totally 266 selected publications are arranged as resource references. 3 Results Six hundred and thirty two taxa have been recorded since 1892, of these 239 species and sub-species are of Crustacea, 368 species and sub-species are of Insecta, 25 species and sub-species are of Arachnida as well as few unidentified species (Table 1). IAEES www.iaees.org 144 Arthropods, 2019, 9(4): 143-175 The presentation format of the tabulated list (Table 1) is as follows: Valid names are given in the first column, together with the indication of the family name; then secondly the first reporter is given with earlier name of species if now changed, thirdly the province(s) name, if from different provinces then all reporters are mentioned.for some species exact place (province name) could not be found, a mark of interrogation is given there. A note on Pakistani freshwater ghost species is also given. Table 1 An inventory of earlier recorded Pakistani species of freshwater free living Arthropoda (Anostraca, Conchostraca, Cladocera, Ostracoda, Copepoda, Peracarida, Arachnida and Insecta). First Reporters from various provinces of Species Province Pakistan Anostraca Thamnocephalidae Qadri and Baqai, 1956 as Strepto cephalus Branchinella (Branchinellites) karachiensis and Branchinella karachiensis fide Sindh maduraiensis Belk and Esparaza, 1995 Qadri and Baqai, 1956 as Strepto cephalus Branchinella hardingi Sindh hardingi Phallocryptus spinosa Brtek and Thiery, 1995 as Branchinella spinosa ? Streptocephalidae Branchipus schaefferi Gurney, 1906 Lync Sindh Streptocephalus dichotomus Kazmi and Sultana, 2014 Baluchistan Ghauri and Mahoon, 1980 Streptocephalus lahorensis nomen Punjab Rogers and Padhye, 2014 dubium fide Rogers and Padhye, 2015 Streptocephalus simplex Tanymastix stagnalis Gurney, 1906 as Branchipus pisciformis; Kemp, 1911as Branchipus stagnalis Sindh Kemp, 1911 Punjab ? Simocephalus exspinosus Daphnia similoides Orlova-Bienkowskaja, 2001 Arora, 1931; Jahangir et al., 2004; Iqbal and Kazmi, 1980 Iqbal and Kazmi, 1980 Brehm, 1914 Daphnia fusca Gurney, 1906; Arora, 1931 Daphnia galeata Daphnia psittacea Daphnia rosea Daphnia longispina Naureen, 1998 unpublished thesis Arora, 1931 Chaudari et al., 1986 Mahar et al., 2008 Baluchistan Baluchistan KP Punjab Punjab Punjab Punjab Sindh Daphnia scholderi Ali, 1973; Iqbal and Kazmi, 1990 Punjab, KP., Baluchistan Daphnia hyalina Kilani and mahoon, 1989 unpublished Biswas, 1971 Punjab Daphnia mendotae Kilani and Mahoon, 1989, unpublished Punjab Daphnia laevis Kilani and Mahoon, 1989, unpublished Punjab Daphnia magna Arora, 1931 Arora, 1931, Ghulam Dastagir, 2014 unpublished thesis as Daphnia pulis Naureen, 1998 unpublished thesis Brehm, 1950; Mahar et al., 2008; Iqbal and Punjab Chirocephalidae Chirocephalus priscus Cladocera Daphniidae Simocephalus mixtus Simocephalus vetulus Daphnia pulex Daphnia longiremis Daphnia lumholtzi IAEES Punjab, Sindh, Baluchistan Punjab, Baluchistan Punjab Punjab; Sindh www.iaees.org Arthropods, 2019, 9(4): 143-175 145 Kazmi, 1990 Baluchistan Daphnia straus Mahoon and Sabir, 1985 Punjab Daphnia similis Ceriodaphnia reticulata Mahoon and Sultana, 1977 Mahar et al., 2008; Iqbal and Kazmi, 1990 Punjab Sindh; Bluchistan Ceriodaphnia quadrangulata Kilani and Mmahoon, 1985 unpublished Punjab Ceriodaphnia richardi Kilani and Mmahoon, 1985 unpublished Punjab Ceriodaphnia rotunda Mahoon and Sabir, 1985 as Ceriodaphnia malcay Ceriodaphnia cornuta Jahangir et al., 2000 Sindh Punjab Ceriodaphnia pulchella Baloch et al., 2005 Iqbal and Kazmi, 1990 KP Ceriodaphnia acanthina Iqbal and Kazmi, 1990 Mahoon et al., 1985 Baluchistan Ceriodaphnia megops Mahoon and sabir, 1985; Nawaz et al., 1981 Punjab, KP Ceriodaphnia dubia Nawaz et al., 1981, mahoon and sabir, 1985 as ceriodaphnia meglaops KP, Punjab Ceriodaphnia laticaudata Kilani and mahoon, 1989 unpublished Punjab, Ceriodaphnia regaudi Ceriodaphnia setosa Arora, 1931, Nawaz et al., 1981 Mahar et al., 2014 Punjab, KP Sindh Scapholeberis aurita Sididae Diaphanosoma brachyurum Diaphanosoma sarsi Mahoon and Sultana, 1977 Punjab Iqbal and Kazmi, 1990; Jahangir et al., 2000 Mahar, et al., 2014 Baluchistan; Sindh Sindh Diaphanosoma excisum Iqbal and Kazmi, 1990 Baluchistan Diaphanosoma leuchtenbergianums Kailani and Mahoon, 1987 Punjab Sida crystallina ? Punjab Latonopsis occidentalis Mahoon et al., 1985 not valid Punjab Latonopsis fasciculata Mahoon and Sabir, 1985 Punjab Mahoon and Sabir, 1985 Punjab Macrothricidae Macrothrix laticornis Arora, 1931; Baloch et al., 2000 Punjab; Sindh Macrothrix borysthenica Kailni and Mahoon, 1989 unpublished Pseudosida herricki Macrothrix hirsuticornis Kailni and Mahoon, 1989 unpublished Punjab Punjab Punjab Macrothrix montana Mahoon and Butt, 1985 Macrocthrix rosea Korai et al., 2008; Ghulam Dastagir, 2014 unpublished thesis Sindh, baluchistan Mahoon and Sultana, 1977 Punjab Mahoon, Ghauri and Butt, 1986 Punjab . Wlassicsia kinistinensis Lathonura ovalis nomen inq Bosminidae IAEES www.iaees.org 146 Arthropods, 2019, 9(4): 143-175 Bosminopsis deitersi Bosmina longirostris Jahangir et al., 2000 Baloch et al., 2005; Iqbal and Kazmi, 1980 Sindh Punjab, Baluchistan Bosmina coregoni Chydoridae Alonacostata Alona guttata Mahar et al., 2010, Iqbal and Kazmi, 1980 Sindh, Baluchistan Mahoon and Sultana, 1977 Punjab Alona intermedia Alona longispina=Alonapulchellacomplex Alona quadrangularis Kailani and Mahoon, 1989 unpublished Mahoon and Sultana, 1977 Alonella excisa Kailani and Mahoon, 1989 unpublished Alonella dentifera Kailani and Mahoon, 1989 unpublished Alonella aureola Kailani and Mahoon, 1989 unpublished Karualona karua Kailani and Mahoon, 1989 unpublished Chydorus faviformis Kailani and Mahoon, 1989 unpublished Chydorus gibbus Kailani and Mahoon, 1989 unpublished as Alona karua Punjab Korai et al., 2008 Lashari et al., 2014 Korai et al., 2008 Korai et al., 2008 Kailani and Mahoon, 1989 unpublished, Jafri et al., 1999; Iqbal and Kazmi, 1990 Iqbal and Kazmi, 1990 Kailani and Mahoon, 1989 unpublished Kailani and Mahoon, 1989 unpublished as Alona monacantha Kailani and Mahoon, 1989 unpublished as Kurzialatissima Kailani and Mahoon, 1989 unpublished as Alonella diaphana Mahar et al., 2008 Kailani and Mahoon, 1989 unpublished Kailani and Mahoon, 1989 unpublished Arora, 1931 Kailani and Mahoon, 1989 unpublished Arora, 1931 Kailani and Mahoon, 1989 unpublished Kailani and Mahoon, 1989 unpublished Kailani and Mahoon, 1989 unpublished as Leydigia quadrangularis Mishkatullah and Mahmood, 2014 Arora, 1931; Kailani and Mahoon, 1989 unpublished as Pleuroxus hastatus Kailani and Mahoon, 1989 unpublished Arora, 1931 Kailani and Mahoon, 1989 unpublished Nawaz et al., 1981; Mahoon and butt, 1985 as Chyrodorus globossus Sindh Sindh Sindh Sindh Punjab, Sindh, Baluchistan Jafri et al., 1999, Khan et al., 2014 as Chyrodorus poppei KP, Punjab Sindh Arora, 1931 Arora, 1931 ; Punjab Punjab , Chydorus latus Chydorus eurynotus Chydorus eurynotus brehmi Chydorus ovalis Chydorus parvus Chydorus sphaericus Chydorus barroisi Coronatella monacantha Kurzia angusticaudata Leberis diaphanous Dunhevedia crassa Dunhevedia mullahensis Dunhevedia kingi Oxyurella tenuicaudis Oxyurella brooksi Pleuroxus aduncus Pleuroxus trigonellus Graptoleberis testudinaria Leydigia leydigi Leydigia sp Leydigia acanthocercoides Picripleuroxus laevis Picripleuroxus striatus Peudochydorus globosus Ephemeroporus poppei Moinidae Moina macrocopa Moina brachiata IAEES Mahoon and Sultana, 1977; Mahar et al., 2008 Kailani and Mahoon, 1989 unpublished Sindh, Punjab Punjab Punjab Punjab Punjab Punjab Punjab Punjab Baluchistan Punjab Punjab Punjab Punjab. Sindh Punjab Punjab Punjab Punjab Punjab Punjab Punjab Baluchistan Punjab Punjab Punjab Punjab www.iaees.org Arthropods, 2019, 9(4): 143-175 Moina rectirostris Moina micrura Moina affinis Moina elliptica (species inquirenda) Moina chankensis record dubious fide Xian-Fen Xiang et al., 2015 Moina pubescenta Ilyocryptidae Ilyocryptus spinifer Conchostraca Cyzicidae Eocyzicus swatiensis nomen nudum fide Rogers and Padhye, 2015 Eocyzicus afzali nomen nudum fide Rogers and Padhye, 2015 Eocyzicus bouvieri Caenestheria propinqual Notostraca Triopsidae Triops cancriformis Unidentified Triops Triops longicaudatus Laevicaudata Lynceidae Lynceus baylyi Lynceus brachyurus 147 Baloch et al., 2004 Mahoon and Sultana, 1977 Jahangir et al., 2004 Iqbal and Kazmi, 1980; Naureen, 1998 unpublished thesis as Moina irrasa Nawaz et al., 1981, Mahoon et al.1985 Arora, 1931 as Mediomoinaelliptica Mahar, 2008 Sindh Punjab Sindh Baluchistan,Punjab Kailani and Mahoon, 1989 unpublished Punjab Arora, 1931 Punjab Chaudhry et al., 1978 KP Bibi and Mahoon, 1985 Punjab Qadri and Baqai, 1956 as Streptocephalus maliricus Shakoori, 1968 Sindh Shaheen and Mahoon, 1993 Vredenburg, 1905 Siddiqi and Suleman, 1977 as Apus longicaudatus Punjab Baluchistan KP Rogers and Olsen, 2016 Gurney, 1906 ? Gilgit-Bultistan Khan et al., 2014 Punjab Reddy, 2011 Khan et al., 2014 Punjab Punjab Mahar, 2003 unpublished thesis; Baloch, 2004; Ayub et al., 2018 Sindh, Punjab Baluchistan. Punjab Punjab Sindh Punjab Copepoda Centropagidae Osphranticum labronectum Parastenocarididae Parastenocaris sutlej Parastenocaris lacustris Laophontidae Onychocamptus mohammad IAEES www.iaees.org 148 Cyclopidae Acanthocyclops viridis Acanthocyclops brevispinosus Acanthocyclops vernalis vernalis Cyclops strenuous Cyclops juri Cyclops landei Diacyclops bicuspidatus Diacyclops nanus Eucyclops agilis Arthropods, 2019, 9(4): 143-175 Maqbool et al., 2015 Maqbool et al., 2015 Siddiqi and Suleman, 1977 as Cyclops vernalis Maqbool et al., 2015 Parveen, Mahoon and Saleem, 1990 Mahoon and Zia, 1985 Maqbool et al., 2014 Maqbool et al., 2014 Mahoon and Sultana, 1977; Arshad et al., 1978 Chaudari et al., 1986 Mahar et al., 2009 Maqbool et al., 2014 Maqbool et al., 2014 Mahoon and Zia, 1985 Mahoon and Zia, 1985 Mahoon and Zia, 1985 Mahoon and Zia, 1985 Chaudari et al., 1986; Mahar et al., 2009 Maqbool et al., 2014 Maqbool et al., 2014 Maqbool et al., 2014 Baloch et al., 2004; Maqbool et al., 2014; Arshad et al., 1978 Najam-un-Nisa, Mahoon and Irfan Khan, 1987 Punjab Punjab KP Punjab Punjab Punjab Punjab Punjab Punjab, KP Punjab Sindh Punjab Punjab Punjab Punjab Punjab Punjab Punjab, Sindh Punjab Punjab Punjab Sindh; Punjab; KP Mesocyclops forbesi Mesocyclops mahooni Microcyclops rubellus Najam-un-Nisa, Mahoon and Khan, 1987 Bashir, 1998 unpublshed thesis Maqbool et al., 2014 Microcyclops varicans Mahar et al., 2010; Maqbool et al., 2014 Microcyclops bicolor Microcyclopslongiramus Paracyclops fimbriatus Paracyclops multanensis Paracyclops affinis Baig and Khan, 1976 Mahar, 2016 Leghari et al., 2003 Bashir, 1998 unpublished thesis Chaudhari et al., 1986 Gurney, 1933 Baloch et al., 2009 Sewell, 1957 Defaye et al., 1987 Lindberg, 1941 Baloch et al., 2004 Punjab Punjab Punjab Sindh Punjab Sindh Sindh Sindh Punjab Sindh Punjab Sindh Punjab Punjab ? Sindh Fada website, 2010 Baqai and Rehana, 1974 as Diaptomus dorsalis Bashir, 1998 unpublished thesis Bashir, 1998 unpublished thesis Bashir, 1998 unpublished thesis ? Sindh Punjab Punjab Punjab Eucyclop macrurus Eucyclop serrulatus Ectocyclops phaleratus Ectocyclops bradyi Ectocyclops clausi Ectocyclops forbesi Ectocyclops sarsi Macrocyclops albidus Macrocyclops fuscus Mesocyclops aspericornis Mesocyclops edax Mesocyclops leuckarti Mesocyclops cokeri nomen dubium Thermocyclops hyalinus Thermocyclops vermifer Thermocyclops rylovi Thermocyclops tinctus Tropocyclops prasinus Diaptomidae Arctodiaptomus salinus Arctodiaptomus dorsalis Diaptomus ahsanulislami Diaptomus pakistanicus Diaoptomus punjabicuss IAEES Punjab www.iaees.org Arthropods, 2019, 9(4): 143-175 Diaptomus castor Heliodiaptomus viduus Heliodiaptomus cinctus Leptodiaptomus siciloides Neodiaptomus kingherensis Paradiaptomus greeni Skistodiaptomus pallidus Skistodiaptomus pygmaeus Skistodiaptomus oregonesis Eudiaptomusgracilis Ergasilidae Ergasilus genuinus Ostracoda Cyprididae Bradleystrandesia reticulata Cyclocypris globosa Cypria mediana Cypris kumara Cypris matthai Cypris pubera Cypris subglobosa Cypridopsis globulus Cypretta nigra Cypretta turgida Dolerocypris sinensis Eucypris virens Eucypris matthai Herpetocypris fontinalis Heterocypris zugmayeri Stenocypris cylindrical major Physocypria devai Stenocypris fontinalis Cypridopsidae Cypridopsis vidua Cypridopsis dentomarginata Heterocypris incongruens Heterocypris salina Heterocypris chandrai Cyprinotus madyensis Cyprinotus crenatus Cyprinotus inaequivalvis Illyocypris bradyi Potamocypris villosa Potamocypris steueri . IAEES Khan et al., 2014 Mahar et al., 2007; Baqai and Rehana, 1974 as Neodiaptomus kamakhiae; Mahar et al., 2007 Maqbool et al., 2015 Baqai et al., 1975 Gurney, 1916 Maqbool et al., 2014 Maqbool et al., 2015 Maqbool et al., 2014 Arshad et al., 1980 as Diaptomusgracilis Lashari et al., 2014 as Limnoncaea genuine Arora, 1935 Khan et al., 2014 Korai et al., 2014 Arora, 1931 as Eucypris kumari 149 Sindh Sindh Punjab Sindh Punjab Punjab Punjab Punjab KP Sindh Punjab Sindh Punjab Sindh Gurney, 1920 as Eucypris pubera Mahar and Jafri, 2012 Arora, 1931 as Cypretta globulus Arora, 1935 Arshad et al., 1980 Mahar and Jafri, 2012 as D. simensis Mahar and Jafri, 2012 Arora, 1931 as Eurycypris matthai Mahar, 2008 Brehm, 1914 as Cyprinotus zugmayeri Arora, 1931 as Stenocypris malcomsoni Arora, 1935 Arshad et al., 1980 as Stenocypris malcomsoni Arora, 1931 as Cypria devai Mahoon and Sultana, 1977 Baluchistan Sindh Punjab Sindh KP Sindh Sindh Punjab Sindh Baluchistan Punjab Sindh Mahar, 2008 as Cypridopsis obesa Gurney, 1920 as Cypridopsis dentomarginatus Gurney, 1920; Arora, 1931; Arshad et al., 1980 as Cyprinotus incongruens Arora, 1935 as Cyprinotus fretens Chaudhry et al., 1978 Arora, 1931 as Cyprinotus chandrai Arora, 1931 Arora, 1931 Hartman, 1964 Gurney, 1920 Arora, 1931 Sindh Baluchistan Baluchistan; Punjab; KP Sindh KP Punjab Sindh Punjab ? Baluchistan Punjab Gurney, 1920 Baluchistan KP,Punjab Punjab www.iaees.org 150 Limnocytheridae Limnocythere inopinata Amphipoda Anisogammaridae Anisogammarus madyensis Mysida Mysidae Mesopodopsis orientalis Decapoda Palaeomonidae Palaemon styliferus Macrobrachium altifrons ranjhai Arthropods, 2019, 9(4): 143-175 Gurney, 1920 Baluchistan Chaudhri et al., 1978 KP Kazmi and Sultana, 2015 Sindh Shakoor, 1968 Sindh, Baluchistan, Punjab Tiwari, 1963 KP Qureshi, 1956 Shakoor, 1968 Kazmi and Kazmi, 2012 as Macrobrachium dacqueti Qureshi, 1956 Shakoor, 1968 Shakoor, 1968 Sindh, Baluchistan, Punjab Sindh, Punjab Macrobrachium semmelinkii Shakoor, 1968 Shakoor, 1968 Ali, 1973 as Palaemonmalcolmsoni Kazmi and Kazmi, 2012 Baluchistan, Sindh Sindh, KP, Baluchistan Punjab Sindh Baluchistan Macrobrachium naraensi Qadri, 1960 Sindh Baluchistan Macrobrachium shahpuri Kazmi and Kazmi, 2012 Punjab Macrobrachium taunsii Kazmi and Kazmi, 2012 Punjab Macrobrachium tirmiziae Yaqoob and Kazmi, 1987 Punjab Macrobrachium rude Husain, 1973 Sindh Macrobrachium idella Yaqoob, 1986 as Macrobrachium idea Sindh Macrobrachium naso Siddiqui, 1976 Sindh Palaemonetes sp Qadri, 1960 Sindh Caridina weberi sumatrensis Kazmi and Siddiqui, 2002 Sindh Caridina nolitica Mehr et al., 1988 Punjab Macrobrachium rosenbergii Macrobrachium lamarrei Macrobrachium dayanus Macrobrachium scabriculum Macrobrachium malcolmsonii Sindh Sindh, Baluchistan, Punjab Sindh Atyidae Caridina rajadhari http://aquaforum.lviv.ua/forum/index.php ? Caridella sp. Baqai et al., 1974 Sindh Kalriana sunahrensis Kalriana karachi Zuberi, 1990 Zuberi, 1990 Sindh Sindh Kalriana jhimphirensis Zuberi, 1990 Sindh Kalriana anissi Zuberi, 1990 Sindh IAEES www.iaees.org Arthropods, 2019, 9(4): 143-175 151 Penaeidae Parapenaeopsis stylifera Korai et al., 2008 Sindh Penaeus indicus Korai et al., 2008 Sindh Penaeus japonicus Korai et al., 2008 Sindh Penaeus merguiensis Metapenaeus brevicornis Brachyura Gecarcinucidae Maydelliathelphusa masoniana Korai et al., 2008 Korai et al., 2008 Sindh Sindh Henderson, 1893 as Thelphusa masoniana Punjab, KP Sartoriana cf. spinigera Alcock, 1910 as Thelphusa spinigera Sindh Sindh,Punjab, KP Sodhiana afghaniensis Kazmi et al., 2005 as Sartoriana afghaniensis Baluchistan Sodhiana blanfordi Alcock, 1909 as Sartoriana blanfordi Pretzmann, 1967 Baluchistan Oziotelphusa sp Kazmi et al., 2006 Sindh Acanthopotamon martensi Henderson, 1893 as Paratelphusa martensi KP Kanpotamon simulum Chaudhri et al., 1978 as Potamon (P) simulum KP Himalayapotamon emphyseteum Kazmi et al., 2005 Punjab Himalayapotamon koolooense Alcock, 1910 as Potamon koolooense Alcock, 1910 Pretzmann, 1963 Pretzmann, 1965 as Potamon (Orientopotamon) gedrosianum waziristanis Hashmi, 1964 as Potaman (Potamonates) sidneyi Henderson, 1893 as Potamon fluviatie Ali, 1973 KP, Punjab Tirmizi and Ghani, 1996 Sindh Rehana et al., 2000 Punjab Rehana et al., 2000 Punjab Potamidae Potamon gedrosianum Potamonautes perlatum Telphusa fluviatalis KP, Punjab, Baluchistan FATA Sindh Baluchistan Punjab, KP Varunidae Varuna litterata Insecta Bourletiellidae Bourletiella sp Isotomidae Isotomurus sp IAEES www.iaees.org 152 Arthropods, 2019, 9(4): 143-175 Saldidae Micracanthia minor Micracanthia ornatula Macrosaldula jakowleffi Saldula orthochila Saldula xanthochila Saldula burmanica Saldula.palustris Leptopodidae Valleriola cicindeloides Oligoneuridae Oligoneuriella kashmirensis Lachlania sp Leptophlebiidae Choroterpes quadriica Ephemeridae Ephemera annandalei Ephemera brunnea Ephemera soanensis Ephemerellidae Torleya swatensis Torleya nepalica (nymph) Crinitella nasiri Isonychiidae Lsonychia khyberensis Baetidae Baetis macanis Baetis meeheanis Cloeon apicatum Cloeon karachiensis Cloeon gilliesi unjustified emendation ICZN (1999) Art. 33.2.3. Centroptilum sp. Psudocloeon sp Amelitidae Amwletus sp Caenidae Caenis kimminsis Caenis izhari Caenis amini Heptageniidae (= Ecdyonuridae) Ecdyonurus pakistanicus Ecdyonurus hazaraensis Hamid and Sutana, 1972 as Saldulaminor,Vinokurov, 2012 Polhemus and Polhemus, 2012 Vinokurov and Kment, 2015 Vinokurov and Kment, 2015 Chen and Lindskog, 1994 Lindskog, 1975 Hamid and Sutana, 1972 Sindh Gilgit-Baltistan Gilgit-Balistan Punjab Gilgit-Balistan Punjab Hamid, 1971, as Leptopus cicindeloides Sindh Ali, 1971 as Oligoneura kashmirensis Ahsan and Aazizullah, 1974 AJK KP Ali, 1967 as Choroterpes (Euthraulus) quadricus Ali Punjab Chopra in Hafiz, 1937 Ali, 1970a Ali, 1970 b as Ephemera striatus Ali, 1967 as Ephemera (Ephemera) soanica Sindh KP Punjab Punjab Ali, 1971 as Ephemerella swatensis Ali, 1971 as Ephemera wahensis Ali, 1971a as Ephemerella nasiri Ali, 1971b as Torleya nasiri KP Punjab, AJK Punjab,KP AJK Ali, 1970 as Eatonia khyberensis KP Ali, 1967 as Baetis macani Ali, 1967 Navas, 1931 Ali, 1970 Punjab Punjab Baluchistan Sindh Ali, 1967 as Cloeon gillican Punjab Ahsan and Azizullah, 1974 Ahsan and Azizullah, 1974 KP KP Ahsan and Azizullah, 1974 KP Ali, 1967 Ali, 1967 Ali, 1967 Punjab KP KP Braasch, 1984 incerta sedis Ali, 1970 as Heptagenia hazaraensis Punjab KP Notacanthurus islamabadicus (nymph, adult) Ali, 1967, 1973 as Ecdyonurus islamabadensis Punjab Rhithrogena baser Palingeniidae Anagenesia minor Coenagrionidae Ali, 1971 KP Eaton, 1892 Sindh Kanth, 1985 Khaliq and Maula , 1999 Kanth, 1985 Khaliq et al., 1990 Tsuda, 1991 Khaliq, 1990 Khaliq and Siddique, 1995 Khaliq , 1990 Ahmed et al., 2008 Khaliq, 1990 AJK KP AJK, Punjab, Sindh, Baluchistan, KP ? Baluchistan AJK KP AJK Punjab Aciagrion hisopa Agriocnemis pygmaea Agriocnemis splendidissima Agriocnemis dabreui Agriocnemis nana IAEES www.iaees.org Arthropods, 2019, 9(4): 143-175 Amphiagrion sp Ceriagrion coromandelianum Ceriagrion cerinorubellum Ceriagrion pulchellum (naiads) Ceriagrion tenellum (naiads) Coenagrion naiads Enallagma cyathigerum Amphiallagma parvum Helochares anchoralis Ischnura fountainei Ischnura senegalensis Ali, 1973 Morton, 1907 Kanth, 1985 Khaliq, 1990 Ahmed et al., 2009 Kanth , 1985 Khaliq , 1990 Din et al., 2013 Din et al., 2013 Ali, 1973 Khaliq et al., 1994 Kanth , 1985 as Enallagma parvum Khaliq , 1990 as Enallagma parvum Ahmed, 2016 as Enallagma parvum Darilmaz and Ahmed, 2015 Khaliq , 1990 Azad et al., 2016 Khaliq , 1990 Khaliq and Siddique, 1995 Ahmed et al., 2009 Ischnura aurora Khaliq , 1990 Kanth , 1985 Ahmed et al., 2009 Ischnura aurora rubilio Kanth, 1985 Tsuda, 1991; Mitra and Babu , 2009 as Ischnura aurora aurora Khaliq and Maula, 1999 Ahmed, unpublished, Ahmed et al., 2011 Ischnura elegans elegans Ischnura forcipata Fraser , 1919 Kanth , 1985 Ahmed et al.2009 Mitra and Babu, 2009 Morton, 1907; Fraser, 1923 Kanth, 1985 153 Punjab Baluchistan AJK Punjab, KP, Sindh, Gilgit AJK Punjab Punjab Punjab Punjab Gilgit-Baltistan AJK Punjab and KP Gilgit - Baltistan Sindh Baluchistan KP Baluchistan, Punjab, Sindh AJK Gilgit-Baltistan Baluchistan, Punjab, KP, Sindh AJK Gilgit-Baltistan AJK , Punjab ?, Sindh, Baluchistan KP Sindh, Punjab. Gilgit-Baltistan KP, Baluchistan AJK Gilgit-Baltistan KP , Punjab Baluchistan Punjab AJK Punjab, Baluchistan, KP Gilgit-Baltistan Punjab and Sindh Ischnura inarmata Ishnura bhimtalensis Mortonagrion gautama Nehalennia sp Paracymus aeneus Khaliq, 1990 Ahmed et al., 2009 Mitra and Babu , 2009 Khan, 2016 Mitra and Babu , 2009 Khan, 2016 Hussain , 2006 Ali, 1973 Darilmaz and Ahmed, 2015 Pseudagrion decorum Kanth , 1985; Khaliq , 1990 Pseudagrion hypermelas Kanth , 1985; Khaliq , 1990 Pseudagrion laidlawi Fraser , 1933; Kanth, 1985; Khaliq, 1990 Pseudagrion microcephalum Mitra and Babu, 2009 Pseudagrion rubriceps Kanth , 1985; Niazi, 1984; Khaliq , 1990; Ahmed, 2016 AJK Punjab , KP Gilgit-Baltistan Pseudagrion spencei Fraser, 1922; Kanth, 1985; Khaliq , 1990; Ahmed, 2016 Sindh AJK Punjab, KP, Baluchistan Gilgit-Baltistan Ischnura delicata IAEES Punjab, Sindh Punjab Gilgit-Baltistan Punjab Sindh AJK Punjab, Sindh, KP, Baluchistan AJK. Punjab Sindh AJK Punjab, Sindh, KP Sindh www.iaees.org 154 Pseudagrion ceylanicum Arthropods, 2019, 9(4): 143-175 Ahmed, 2011; Azad et al., 2016 AJK KP Agrion aurora Punjab Agrion cerinum Paracercion calamorum dyeri Paracercion malayanum Khan , 2016 Khaliq , 1990 Mitra and Babu, 2009 Morton, 1907 Rhodischnura nursei= Ischnura nursei Fraser, 1933 Sindh Kanth, 1985 AJK Punjab, Sindh, KP, Baluchistan Punjab KP KP, Sindh ? ? Nanosura aurora Khaliq , 1990 Khan, 2000 Regimbartia attenuata Sternolophus rufipes Sternolophus decens Sternolophus solieri Euphaeidae Hansen , 1999 Darilmaz and Ahmed, 2015 Nasserzadeh et al., 2017 Nasserzadeh et al., 2017 Bayadera indica Kanth, 1985; Khaliq, 1990 Bayadera longicauda Ahmed et al., 2011 ;Yousuf et al., 2000 Epallage fatime Lestidae Archilestes sp Lestes patricia Lestes praemorsus praemorsus Schmidt, 1961 Lestes thoracicus Kanth , 1985; Khaliq , 1990 Lestes viridulus Niazi, 1984 ;Rafi et al., 2009 Sympecma paedisca Morton, 1907 as Sympecma paedisca fusca; Laidlaw, 1920 as Sympecma annulata Synlestidae Cylonolestes cyanea Yousuf et al., 2000 Megalestes major Kanth, 1985; Khaliq, 1990; Ahmed et al., 2009 Ali, 1973 Rafi et al., 2009 Yousuf et al., 2000 Punjab Punjab Baluchistan AJK. KP, Punjab Punjab AJK Baluchistan AJK AJK AJK AJK SindhPunjab Punjab AJK Baluchistan AJK AJK AJK Punjab, KP Gilgit-Baltistan Platycnemidae Calicnemis eximia Khaliq, 1990; Khaliq et al., 1990. Calicnemia fortis Dow et al., 2014 Coeliccia renifera. Khaliq, 1990; Ahmed et al., 2008 Coeliccia vacca Copera ciliata Ahmed, 2016 Khaliq, 1990 Copera marginipes. Kanth, 1985; Khaliq, 1990; Ahmed, 2016 Platycnemis dealbata Platystictidae Drepanosticta polychromatica Drepanosticta carmichaeli Nemouridae Mitra and Babu , 2009 Punjab, KP. AJK AJK Punjab AJK Gilgit-Baltistan Punjab AJK Punjab, KP Gilgit Punjab Khaliq , 1990 Ahmed, 2016 Punjab Gilgit-Baltistan Amphinemura tricantha Amphinemura mirabilis Illiesonemoura atripes Illiesonemoura battakundi Illiesonemoura besali Illiesonemoura lilami Illiesonemoura maluksari IAEES Aubert, 1959 as Amphinemura schmidi and Nemoura schmidi Martynov, 1928 Aubert, 1959 as Nemoura (Nemoura) atripes Aubert, 1959 as Nemoura (Nemoura) battakundi Aubert, 1959 as Nemoura besali Aubert, 1959 as Nemoura (Nemoura) lilami Aubert, 1959 as Nemoura (Nemoura) maluksari AJK AJK KP KP KP AJK KP www.iaees.org Arthropods, 2019, 9(4): 143-175 155 Illiesonemoura pakistani Illiesonemoura polystigma Mesonemoura skardui Aubert, 1959 as Nemoura (Nemoura) pakistani Aubert, 1959 as Nemoura (Nemoura) polystigma Aubert, 1959 Mesonemoura vaillanti Navas, 1922 Capniidae Eucapnopsis stigmatica transversa Perlodidae Aubert, 1959 KP Zhiltzovaia cachemirica Aubert, 1959 as Perlodes (Skobeleva) cachemirica KP Isogenus sp Ahsan and Azizullah, 1974 KP Aubert, 1959 AJK, Punjab, KP Zwick, 1980 Aubert, 1959 Ahsan and Azizullah, 1974 KP AJK KP Anaciaeshna jaspidea Anax indicus Ahmed et al., 2009; Chaudry , 2013 unpublished thesis Chaudri et al., 2013 Chaudri et al., 2010 Anax immaculifrons Fraser, 1936; Rafi et al., 2009; Chaudri, 2010 Anax nigrofasciatus Chaudri, 2010; Chaudri et al., 2016 Anax parthenope Rafi et al., 2009; Chaudri, 2010 unpublished Anax nigrolineatus Anax imperator imperator Boyeria irene Cephalaeschna masoni Gynacanthaeshna sikkima Gynacantha apicalis Hemianax ephippiger Polycanthagyna erythromelas Cordulegastridae Rafi et al., 2009 Fraser, 1936 Din et al., 2013 Akhtar et al., 2014 Chaudri et al., 2010 Fraser, 1927, 1934b Din et al., 2013 Conniff, 2016 Gilgit-Baltistan KP Sindh Punjab KP AJK Punjab Gilgit-Baltistan KP AJK KP, Punjab, Sindh, Baluchistan, Gilgit AJK KP Punjab KP Punjab, KP Punjab Punjab Gilgit-Baltistan Cordulegaster brevistigma brevistigma Yousuf , 1972 unpublished as Kuldanagaste r pakistanica Tsuda, 1991; Chaudri, 2016; Mehmood, 2014 ? Punjab KP , Baltistan AJK Anormogomphus kiritschenkoi Fraser, 1934; Ahmad, 1994 Punjab, Sindh, Bluchistan KP Anormogomphus exilocorpus Anormogomphus heteropterus Anisogomphus vulvalis Yousuf , 1972 unpblished Prasad and Arshney, 1988 Yousuf , 1972 unpublished Burmagomphus pyramidalis Fraser, 1934;Chaudri, 2010 Burmagomphus sivalikensis Erpetogomphus sp Moore (compiler), 1997 Ali, 1973 Chaudri, 2010 as Gomphidia t-nigrum ;Mehmood et al., 2016 Perlidae Neoperla schmidi Chloroperlidae Xanthoperla acuta Xanthoperla kishanganga Alloperla sp. Aeshnidae Aeshna juncea ? AJK Gilgit-Baltistan KP Gomphidae Gomphidia nigrum Ictinogomphus angulosus Chaudri, 2010; Chaudhry , 2016 Ictinogomphus aloquopterus Ictinogomphus pugnovittatus Ictinogomphus rapax Yousuf, 1972 unpublished Yousuf , 1972 unpublished Yousuf and Yunus, 1976 Chaudri, 2010 Chaudri, 2010;Chaudri et al., 2016 ;Mehmud et al, Lamelligomphus biforceps IAEES Punjab ? KP AJK Punjab ? Punjab Punjab KP AJK KP Punjab Punjab Punjab Sindh , AJK, Punjab AJK www.iaees.org 156 Arthropods, 2019, 9(4): 143-175 2016 as Onychogomphus biforceps Ocotgomphius sp Nepogomphus modestus Onychogomphus bistrigatus Ophiogomphus reductus Ophiogomphus caudoforcipus Platygomphus dolabratus Paragomphus lineatus Lindenia tetraphylla Libellulidae Ali, 1973 Chaudri et al., 2016 Perveen et al., 2014 Khaliq et al., 1994 Yousuf, 1972 unpublished Mehmud et al., 2016 Rafi et al., 2009 as Mesogomphus lineatus Rehman et al., 2015 Waterson, 1980 Punjab KP Punjab Punjab KP Gilgit and Baltistan, AJK KP KP AJK Sindh Diplacodes lefebvrei Ahmed et al., 2009; Perveen et al., 2014 ; Ullah et al., 2000 Diplacodes trivialis Perveen et al., 2014 Ullah et al.2000 Punjab, Sindh, Baluchistan, KP, Baltistan Punjab Punjab, Sindh, AJK, KP Punjab, Sindh, KP, Baluchistan, AJK Punjab Sindh Punjab KP Punjab Sindh Punjab Gilgit-Baltistan AJK , Punjab KP Sindh Gilgit-Baltistan KP Sindh KP Sindh Erythemis sp Ali, 1973 Kp Hydrobasileus croceus Leucorrhinia sp Libellula fulva Saeed et al., 2016 Ali, 1973 Libellula quadrimaculata Macromia sp Nannophya pygmaea Nannothemis sp Neurothemis fluctuans Neurothemis tullia tullia Orthemis sp Khaliq et al., 1994 Ali, 1973 Fraser, 1921 Ali, 1973 Chaudri et al., 2016 Din et al., 2013 naiads Lakhiar and Panhwar , 2015 Ali, 1973 Orthetrum anceps Jehangir etal , 2016 Orthetrum brunneum brunneum Fraser, 1936; Ahmed et al., 2009 Orthetrum cancellatum Khaliq et al., 1994 Orthetrum chrysis Din et al., 2013 naiads; Akhtar et al., 2014 KP Punjab Punjab KP Gilgit-Baltistan Punjab KP Punjab Punjab Punjab Sindh Punjab KP Baluchistan ? Gilgit-Baltistan Gilgit-Baltistan Punjab KP Orthetrum luzonicum ( naiads) Din et al., 2013 Orthetrum glaucum Ahmed et al., 2009; Rafi et al., 2009 ; Khan , 2016 naiad Orthetrum japonicum internum Orthetrum chrysostigma luzonicum Tsuda, 1991 Ahmed et al., 2009 Orthetrum pruinosum neglectum Perveen et al., 2014; Rehman et al., 2015 Acisoma panorpoides panorpoides Chaudri, 2010 Belonia sp Brachydiplax sobrina Ali, 1973 Chaudri et al., 2010 Brachythemis contaminata Chaudri2010 Bradinopyga geminate Brechmorhoga sp Crocothemis erythraea Crocothemis servilia IAEES Chaudri, 2010 Rehmani et al., 2015 Ali, 1973 Rafi et al., 2009 Din et al., 2013 naiads Ullah et al., 2000 Kirby , 1886 as Crocothemis reticulata Ahmed et al., 2009; Rafi et al., 2009; Akhtar et al., 2014; Rehman et al., 2015 Din et al., 2013 naiads; Akhtar et al., 2014 Punjab Gilgit-Baltistan KP Punjab ? Gilgit – Baltistan KP Sindh www.iaees.org Arthropods, 2019, 9(4): 143-175 157 Selysiothemis nigra (naiads ) Somatochlora sp Sympetrum naiad Tarnetrum sp. Tauriphila sp. Tramea virginia Tramea basilaris burmeisteri Din et al., 2013 Ali, 1973 Hussain and Ahmed, 2004 Ali, 1973 Ali, 1973 Chaudri et al., 2016 Trithemis annulata insectoid.info Trithemis kirbyi kirbyi Din et al., 2013 naiads; Akhtar et al., 2014 ; Chaudri et al., 2016 Trithemis inervis Rafi et al., 2009 ‘ Din et al., 2013 naiads; Akhtar et al., 2014; Chaudri et al., 2016 Trithemis parviaurora Yousuf, 1972 Punjab KP Gilgit-Baltistan Punjab KP Punjab KP Gilgit-Baltistan AJK KP AJK Punjab , Sindh KP Sindh KP KP Punjab Punjab, Sindh Sindh Punjab Punjab Sindh Punjab Punjab Punjab , KP Punjab , Sindh AJK ? Punjab KP Punjab, Gilgit-Baltistan AJK Punjab KP Sindh ? Trithemis pallidinervis Tholymis tillarga Urothemis signata signata Zygonyx torridus isis Zyxomma petiolatum Chlorocyphidae Libellago greeni Libellago lineata lineata Ullah et al., 2000; Rafi et al., 2009 Chaudri et al., 2016; Fazllulah et al., 2016 Chaudri , 2016 Fraser, 1934; Chaudri, 2016 Chaudri , 2016 Sindh, AJK Sindh, KP Sindh, Punjab, KP Punjab, , P Punjab, AJK Ahmed et al., 2009 Chaudril, 2010 Ali, 1973 Gilgit-Baltistan Punjab, KP, AJK Punjab Ahmed et al., 2011 Ahmed et al., 2011 Ahmed , 2010 unpublished , Ahmed ., 2016 Kanth , 1985 Niazi, 1984 Khaliq 1990 Mitra and Babu, 2009 AJK AJK, KP, Punjab AJK, KP AJK, Punjab, Punjab and KP Punjab Kanth, 1985 Khaliq and Maula, 1999 Kanth , 1985 , Niazi, 1984, Zada et al., 2016 Khaliq et al., 1990 Khaliq et al., 1990 Khaliq , 1990 Khaliq et al. 1990 Khaliq et al., 1990 AJK KP AJK, Punjab, KP AJK AJK KP, Punjab AJK KP, Punjab Orthetrum sabina Din et al., 2013 naiads ; Akhtar et al., 2014 Orthetrum taeniolatum Ahmed et al., 2009; Din et al., 2013 naiads; Akhtar et al., 2014 Orthetrum testaceum testaceum Orthetrum triangulare triangulare Chaudri et al., 2016 Perveen et al., 2014 Palpopleura sexmaculata sexmaculata Ahmed et al., 2009 ; Rafi et al., 2009; Jehangir et al., 2016 Pantala flavescens Rafi et al., 2009; Amad Ud Din et al, 2013 ; Perveen et al., 2014; Rehman et al., 2015 Potamarcha obscura Rhodothemis rufa Saeed et al., 2016 Perveen et al., 2014 Rhyothemis variegata variegata Hussain and Niazi, 1999 naiad ;Yousuf et al., 2013 naiads; Rehman et al., 2015 Protoneuridae Elattoneura atkinsoni Elattoneura souteri Elattoneura campioni Elattoneura nigerrima Elattoneura tetrica Calopterygidae Neurobasis chinensis chinensi Rhinocypha quadrimaculata Rhinocypha immaculata Rhinocypha hilaryae Rhinocypha trifasciata Rhinocypha unimaculata IAEES Yousuf et al., 2013 naiads; Rafi et al., 2009 www.iaees.org 158 Gyrinidae Dineutus indicus Arthropods, 2019, 9(4): 143-175 Ahmed et al., 2008 AJK Mazzoldi, 2003 Hájek and Fery, 2004 Baltistan Orectochilus afghanus Dineutus sp Dineutus spinosus Gyretes sp Patrus haemorrhous Dytiscidae Agabus solskii Agabus biguttatus Agabus conspersus Agabus bipustulatus Agabus debilipes Canthydrus laetabilis Clypeodytes orissaensis Copelatus freudei Rehana et al., 2000 Darilmaz and Ahmed, 2015 Ali, 1973 Darilmaz and Ahmed, 2015 Angelini, 1978 Ghosh and Nilsson , 2012 Ghosh and Nilsson , 2012 Brancucci , 1979 Ghosh and Nilsson , 2012 Guignot, 1958 as Agabus skarduensis Vazirani, 1969 Nilsson, 2012 Vazirani, 1977 Darilmaz and Ahmed, 2015 Cybister tripunctatus lateralis Vazirani , 1969 as Cybister tripunctatus asiaticus Darilmaz and Ahmed, 2009 Cybister cardoni Vazirani, 1969 Vazirani, 1977 Ali, 1973 Vazirani, 1977 Vazirani, 1977 Hajek, 2006 Fery and Hosseinie, 1998 Hájek and Fery, 2004 Hajek, 2006 Miller, 2002 Darilmaz and Ahmed, 2009 Hasan et al., 2013 Guéorguiev , 1967 Miller , 2002 Guéorguiev , 1967 Vazirani , 1972 Guignot, 1959 as Hyphydrus lindemannae Bistrim,1982 Ghosh and Hedge , 2015 Vazirani, 1970 Jach, 2003 Jach, 2003 Jach, 2003 Jach, 2003 Cybister confusus Cybister limbatus Cybister sugillatus Deronectes hendrichi Deronectes bameuli Deronectes afghanicus Eretes griseus Dytiscus adult Eretes sticticus Herophydrus musicus Hyphydrus lyratus flavicans Hyphydrus gschwendtneri Hyphydrus renardi Hygrotus (Coelambus)enneagrammus Hygrotus(Coelambus)confluens Hygrotus(Coelambus) flaviventris Hygrotus(Leptolambus) impressopunctatus Hygrotus (Leptolambus) zigetangco Hygrotus (Hyphoporus) anitae Hygrotus (Hyphoporus )aper Hygrotus (Hyphoporus )severini Hygrotus (Hyphoporus) bertrandi Hygrotus (Hyphoporus )elevatus Hygrotus (Hyphoporus )nilghiricus Hygrotus (Hyphoporus ) pacistanus Hydaticus histrio Hydaticuspictus Hydaticus vittatus vittatus Hydaticus ponticus IAEES Jach, 2003 Vazirani, 1969 as Hyphoporus anitae Hájek, 2006 as Hyphoporus aper Vazirani, 1977 as Hyphoporus severini Vazirani, 1969 as Hyphoporus bertrandi Vazirani, 1977 as Hyphoporus elevatus Vazirani, 1977 as Hyphoporus nilghiricus Guignot, 1959 as Hyphoporus pacistanus Guéorguiev , 1967 Franciscolo, 1968 Angelini, 1978 Vazirani 1977 as Prodaticus pictus Vazirani , 1977 Wewalka, 1979 Darilmaz and Ahmed, 2009 as Hydaticus leander KP ,at the border with Afghanistan Punjab Sindh KP Sindh Baltistan KP, Punjab KP, Punjab KP Punjab Gilgit-Baltistan ? Punjab ? Sindh Baluchistan Sindh ? ? KP ? ? Baluchistan KP KP Baluchistan ? Sindh KP Baluchistan ? Baluchistan ? Punjab ? Baluchistan Baltistan KP KP KP AJK Punjab, Baluchistan Baluchistan ? Punjab ? ? Punjab Baluchistan ? Baltistan Baluchistan ? ? Sindh www.iaees.org Arthropods, 2019, 9(4): 143-175 Hydaticus ricinus Hydaticus fabricii Hydaticus vhantoides Hydronebrius mattheyi mattheyi Hydroglyphus pendjabensis Hydroglyphus signatellus Megadytes sp. Hydroglyphus angularis Hydroglyphus geminus Hydrovatus fusculus Hydrovatus confertus Ilybiosoma kermanensis Laccophilus maindroni persicus Laccophilus minutus Laccophilus indicus Laccophilusinefficiens Laccophilus sharpi Laccophilus flexuosus Laccophilus parvuIus Macrovatellus sp Nebrioporus airumlus Nebrioporus indicus Neptosternus circumductus Peschetius quadricostatus Platynectes ( Gueorguievtes) kashmiranus kashmiranis Platambus lineatus Platambus lindbergi Platambus sogdianus Platambus wewalkai Potamonectes insignis Pseuduvarus vitticollis Rhantus sikkimensis Rhantus taprobanicus Sandracottus dejeani Sandracottus festivus Thermonectus sp Uvarus genitalis Nepidae Nepa ruber Nepa elongates Laccotrephes ruber Cercotmetus asiaticus Cercotmetus pilipes Ranatra filiformis Ranatra digitata Ranatra elongata Corixidae Agraptocorixa hyalinipennis Corixa substriata IAEES Guignot, 1959 Wewalka , 1979 Vazirani, 1969 Wewalka, 1979 Brancucci, 1980 Guignot, 1954 as Guignotus pendjabensis Vazirani , 1969 as Guignotus signatellus Darilmaz and Ahmed, 2015 Ahsan and Azizullah, 1974 Guéorguiev , 1967 as Guignotus angularis Vazirani, 1977 Erman and Erman, 2008 Wewalka, 1982 Wewalka, 1982 Hajek, 2006 Brancucci, 1983 Vazirani , 1969 Vazirani, 1969 as Laccophilus sindensis Hajek, 2006 Vazirani, 1970 as Laccophilus chinensis inefficiens Hajek, 2006 Hajek, 2006 Gschwendtner, 1936 as Laccophilus solutus indicus Guignot, 1959 Fery and Darvishzadeh, 2012 Hajek, 2006 Rehana et al., 2000 Vazirani, 1970 as Potamonectes kashmirensis Angelini , 1978 as Hydrophoruskashmirensis Toledo, 1998 Guignot, 1959 as Potamonectes indicus;Vazirani, 1970 as Potamonectes manii Angelini, 1978 as Nebrioporus manii, Potamonectes manii Hajek, 2006 (likely) Biström and Nilsson, 2003 159 KP ? ? KP Punjab , KP Punjab Baluchistan Sindh KP KP, Baluchistan ? KP ? Baluchistan Baluchistan Baluchistan Sindh Baluchistan ? Baluchistan Baluchistan Punjab Baluchistan ? Punjab Baltistan AJK Gilgit-Baltistan Punjab Gilgit- Baltistan, AJK Wewalka , 1975 Vazirani, 1965 as Platambus guignoti Wewalka, 1975 as Platambus holzschuhi Plazi , 2017 Vazirani, 1977 Vazirani, 1977 Vazirani, 1970 as Rhantuspunjabensisn Brancucci, 1979 Vazirni, 1972 Ghosh and Hedge , 2015 Steven, 2013 Ali, 1973 as Thermonectes sp Vazirani, 1969 Baluchistan Baluchistan AJK,KPat the border with Afghanistan ? Punjab KP KP Baluchistan ? Punjab ? ? Baluchistan KP KP ? Irfan and Bibi, 1999 Irfan and Bibi, 1999 Srivastava, 2008 Cheng et al., 2006 Irfan and Bibi, 1999 Srivastava, 2008 Vazirani , 1970 Ali, 1973 Punjab Punjab ? ? Punjab ? ? KP Thirumalai and Kumar, 2005 Irfan and Bibi, 1999 ? Punjab Nilsson, 1998 www.iaees.org 160 Corixa hieroglyphica Micronecta proba Micronecta thyesta Gerridae Gerris fossarum Neogerris parvula Limnogonus fossarum fossarum Limnogonus sp Veliidae Microvelia diluta Pseudovelia sexualis Halovelia sp. Velia (Plesiovelia) affinis marussii Diplonychus annulatus Belostomatidae Lethocerus indicus Lethocerus indicum Lethocerus patruelis Simuliidae Gymnopais sp Parasimulumsp Prosimuliumsp Hydrophilidae Berosus nigriceps Berosus chinensis Berosus fairmairei Berosus indicus Berosus indiges Coelostoma stultum Enochrus (Methydrus) esuriens Enochrus(Lumetus) ater Enochrus (Lumetus) sinuatus Hydrophilus senegalensis Helochares anchoralis Paracymus aeneus Regimbartia attenuata Sternolophus rufipes Sternolophus decens Sternolophus solieri Spercheidae Spercheus belli belli Notonectidae Anisops cavifrons Anisops sardeus sardeus Notonecta viridis Notonecta maculata Helophoridae Helophorus (Eutrichelophorus) micans Psychomyiidae Polycentropus sp Rhagionidae Atherixsp Chironomidae Prodiamesa sp Polypedilum sp Pentaneura sp Midges Naucoridae IAEES Arthropods, 2019, 9(4): 143-175 Ali, 1973 Irfan and Bibi, 1999 Irfan and Bibi, 1999 Punjab, KP Punjab Punjab Irfan and Bibi, 1999 Thirumalai and Kumar, 2005 Kazmi, in press Hsan and Azizullah , 1974 Punjab ? Sindh KP Irfan and Bibi, 1999 Hecher and Zettel, 2006 Kazmi , in press Andersen, 1995 Jhamalar and Chandra, 2012 Punjab ? Sindh Ali, 1973 as Belostomaindica Irfan and Bibi, 1999 Polhemus, 1995 Punjab, KP Punjab ? Rehana et al., 2000 Rehana et al., 2000 Rehana et al., 2000 Punjab Punjab Punjab Schödl , 1994; Darilmaz and Ahmed, 2015 Hansen , 1999 Hansen, 1999 Hansen, 1999 Darilmaz and Ahmed, 2015 Hansen , 1999 Darilmaz and Ahmed, 2015 Darilmaz and Ahmed, 2015 Hansen , 1999 Darilmaz and Ahmed, 2015 Darilmaz and Ahmed, 2015 Schödl , 1998 Darilmaz and Ahmed, 2009 Darilmaz and Ahmed, 2015 Darilmaz and Ahmed, 2015 Hansen , 1999 Darilmaz and Ahmed, 2015 Nasserzadeh et al., 2017 Nasserzadeh et al., 2017 ? Baltistan ? KP Sindh KP Sindh KP Baluchistan KP, Sindh Sindh ? Sindh Sindh Sindh KP KP, Sindh ? ? Hansen, 1999 Darilmaz and Ahmed, 2015 KP Thirumalai and Kumar, 2005 Irfan and Bibi, 1999 as Anisops sardea; Dash, 2014 Hoberlandt , 1961 Berchi, 2013 ? Punjab ? ? Mart and Erman , 2000 ? Ali, 1973 Punjab Ali, 1973 as Antherix sp in Chironomidae KP Ali, 1973 Ali, 1973 Ali, 1973 Siddiqi and Suleman, 1977 KP KP KP Gilgit Baltistan www.iaees.org Arthropods, 2019, 9(4): 143-175 161 Ambrysus sp. ?Cryphocricas sp Noteridae Canthydrus laetabilis Hydrocoptus subvittulus Pronoterus sp Pleidae Plea sp. Psephenidae Psephenus sp Hydropsychidae Cheumatopsyche sp Smicridea sp Irfan and Bibi, 1999 Ali, 1973 Punjab Sindh Fery, 2012 Wewalka, 1982 Ali, 1973 Baluchistan ? KP Ali, 1973 KP Ali, 1973 Punjab, AJK Ali, 1973 , Ahsan and Azizullah, 1974 Ali, 1973 Punjab, KP Punjab, KP Macronemum sp. Ahsan and Azizullah, 1974 KP Ahsan and Azizullah, 1974 KP Ali, 1973 Punjab Ali, 1973 Punjab Hasan et al., 2013 KP Khatoon and Ali, 1989 Punjab Ahsan and Azizullah, 1974 KP Ahsan and Azizullah, 1974 KP Agapetus sp. Ahsan and Azizullah, 1974 KP Glossosoma sp. Stenopsychidae Stenopsyche sp. Limnephilidae Pseudostenophylax sp Corydalidae Corydalus sp. Blephariceridae Bibiocephala sp. Tipulidae Limoniasp Sialidae Sialis sp. Limnichidae Byrrhinus marginatus Pionidae Hydrochoreutes sp. Homocaligidae Annerossella sp. Blattisociidae Platyseius sp. Arrenuridae Arrenurus madaraszi Ahsan and Azizullah, 1974 KP Ahsan and Azizullah, 1974 KP Ahsan and Azizullah, 1974 KP Ahsan and Azizullah, 1974 KP Ahsan and Azizullah, 1974 KP Rehana et al., 2014 KP Rehana et al., 2014 ? Reisenand Mullen, 1978 Punjab Reisenand Mullen, 1978 Punjab Reisenand Mullen, 1978 Punjab Reisenand Mullen, 1978 Punjab Khatoon and Ali, 1989 Punjab, KP Hydropsyche sp. Polycentropodidae Neureclipsis sp Leptoceridae Mystacides sp Arachnida Araneae Cybaeidae Argyroneta aquatica Acari Limnesiidae Limnesia (Limnesia) lembangensis Psychomyiidae Tinodes sp Dipseudopsidae Phylocentropus sp. Glossosomatidae Hydrachnidae IAEES www.iaees.org 162 Arthropods, 2019, 9(4): 143-175 Hydrachna (Scutohydrachna) testudinata Hydrodromidae Hydrodroma monticola Protziidae Protzia (Calonyx) flagellum Eylaidae Eylais (Metaeylais) hamata Eylais (Protoeylais) degenerata Arrenuridae Arrenurus (Rhinophoracarus) truncatus Lebertiidae Khatoon and Ali, 1989 Punjab Khatoon and Ali, 1989 Punjab Khatoon and Ali, 1989 Punjab Khatoon and Ali, 1989 Khatoon and Ali, 1989 Punjab Punjab Khatoon and Ali, 1989 Punjab, KP Lebertia (Pilolebertia) carmamaya Khatoon and Ali, 1989 Punjab, KP Torrenticolidae Torrenticola (Torrenticola) tetraporella Khatoon and Ali, 1989 Punjab, KP Our survey indicates that the fresh water fauna in Pakistan is shrinking. There are programmes running in the country to conserve and sustainably manage freshwater habitats. Hopefully these freshwater programmes will strive to maintain ecological health, hydrological processes and biological diversity. 4 Discussion 4.1 Pakistani “ghost species” Given in Pakistani literature of fresh water species, some Pakistani “ghost species” whose identities have been shrouded in uncertainty are given below. (A) Palaemonetes sp There are probabilities that Qadri (1960) has mistaken population of Macrobrachium for Palaemonetes as juveniles of Macrobrachium can easily be mistaken for Palaemonetes since they are of same size and similarly found in shallow vegetated areas along the water edge (Kazmi and Kazmi, 2012). (B) Crayfish Akhtar (1995) in his country status paper mentioned of presence of crayfish in KP, Pakistan. Crayfish are freshwater crustaceans resembling small lobsters, to which they are related. Some species are found in brooks and streams where there is running fresh water, while others thrive in swamps, ditches, and paddy fields. Taxonomically, they are members of the superfamilies Astacoidea and Parastacoidea. The greatest diversity of crayfish species is found in south eastern North America, whereas large area of Asia including Pakistan lacks a native crayfish fauna except eastern corner of the continent have no clue what fresh water decapod crustacean Akhtar (1995) took for crayfish. (C) Daphnia Tabassun et al. (2014) mentioned of marine Daphnia which seems to be wrong generic identification. It may be pointed out here that the Cladocera are ubiquitous in inland aquatic habitats, but rare in the oceans. Tabassum et al. (2014) specimens may be some podonids but not Daphnia. (D) Caridella sp- status of Caridella from Pakistan is uncertain. 4.2 Euryhaline genera There are some euryhaline genera- These marine decapods may be discovered in the river system near river mouths in estuarine environments such as Charybdis, Scylla, Varuna and some Penaeid genera. 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Muqtadra Qaumi Zaban, Islamabad, Pakistan Zwick P. 1980. Beitrage zur Kenntnis der Plecoptera des Himalaya. Entomologica Basiliensia, 5: 59-138 IAEES www.iaees.org Arthropods Arthropods play the role of both pests and beneficial organisms. Some arthropods are important crop pests but others are natural enemies. Some arthropods are important health pests but many crustaceans are important food sources of humankinds. Arthropods govern the structures and functions of natural ecosystems, but are always ignored by researchers. On the global scale, the surveys of mammals, birds and vascular plants were relatively perfect because they were economically important and easily surveyed. However, arthropods, despite their ecological and economical importance, have not yet been fully surveyed and recorded due to their difficulties to be sampled. The research on arthropods must be further promoted. The journal, Arthropods, aims to provide a public and appropriate platform for the publication of studies and reports on arthropods. Arthropods (ISSN 2224-4255) is an international open access (BOAI definition), open peer reviewed online journal (users are free to read, download, copy, distribute, print, search, or link to the full texts of the articles) devoted to the publication of articles on various aspects of arthropods, e.g., ecology, biogeography, systematics, biodiversity (species diversity, genetic diversity, et al.), conservation, molecular biology, biochemistry, physiology, control, etc. The journal provides a forum for examining the importance of arthropods in biosphere (both terrestrial and marine ecosystems) and human life in such fields as agriculture, forestry, fishery, environmental management and human health. The scope of Arthropods is wide and includes all arthropods-insects, arachnids, crustaceans, centipedes, millipedes, and other arthropods. Articles/short communications on new taxa (species, genus, families, orders, etc.) of arthropods are particularly welcome. Authors can submit their works to the email box of this journal, arthropods@iaees.org. All manuscripts submitted to Arthropods must be previously unpublished and may not be considered for publication elsewhere at any time during review period of this journal. In addition to free submissions from authors around the world, special issues are also accepted. The organizer of a special issue can collect submissions (yielded from a research project, a research group, etc.) on a specific topic, or submissions of a conference for publication of special issue. Editorial Office: arthropods@iaees.org Publisher: International Academy of Ecology and Environmental Sciences Address: Unit 3, 6/F., Kam Hon Industrial Building, 8 Wang Kwun Road, Kowloon Bay, Hong Kong E-mail: office@iaees.org Arthropods ISSN 2224-4255 Volume 8, Number 4, 1 December 2019 Articles Walking leg regeneration observed in three families and four species of antarctic sea spiders John A. Fornshell 110-117 First record of the crab, Droippe quadridens (Fabricius, 1793) (Brachyura: Dorippidae), from the Iraqi coastal waters of the NW Arabian Gulf, with notes on the occurrence of seven species of crabs in the region KhaledKh Al-Khafaji, Tariq H.Y. Al-Maliky, et al. 118-126 Xylophagous millipede surface area to volume ratios are size-dependent in forests Mark Cooper 127-136 Size dimorphism in six juliform millipedes Mark Cooper 137-142 Inventory of freshwater arthropods in Pakistan Quddusi B. Kazmi, Farhana S. Ghory 143-175 IAEES http://www.iaees.org/