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Land-locked anchialine blue holes are karstic sinkholes and caves with tidally influenced, vertically stratified water bodies that harbor endemic fauna exhibiting variable troglomorphic features. These habitats represent island-like systems, which can serve to elucidate evolutionary and biogeographic processes at local scales. We investigated whether the ‘continuous spelean corridor’ hypothesis may elucidate the biogeographical distributions of the stygobitic annelid Pelagomacellicephala iliffei (Polynoidae) collected from the Great Bahama and Caicos Banks of the Bahamas Archipelago. Phylogenetic reconstructions were performed using Bayesian Inference on individual and combined datasets of three molecular markers (16S rDNA, COI, 18S rDNA) and species delimitation employed three widely accepted methods in DNA taxonomy, namely GMYC, bPTP, and ABGD. Mantel tests were used to test the effect of geography on genetic structure. Using these analyses, we recovered five independently evolving entities of the focal species across four islands of the Great Bahama Bank including Cat, Eleuthera, Exumas, and Long. Genetic data yielded strong correlations between islands and phylogenetic entities, signifying independent evolutionary histories within anchialine caves across the platform. The island of Eleuthera showed intra-island gene flow and dispersal capabilities between blue holes separated by 115 km, providing evidence of a crevicular spelean corridor within the island. However, no evidence of inter-island dispersal is present in the analyzed system. Consistent with previous biogeographic studies of cave crustaceans, the major barriers shaping the cave biota of the Bahamas Archipelago appears to be the deep trenches and channels separating the Bahamian banks.

Malagabdella niarchosorum, new species Figures 8���15 TYPE MATERIAL: Holotype, specimen fixed in 95% ethanol, Madagascar ��� Parc National Ranomafana in Talatakely (selectively logged, low montane forest; MG02-05), March 5, 2002. Collected by Mark Siddall, Elizabeth Borda, Evon Hekkala, Clara Imboule, and Raina Rakotondriany (AMNH 5285, Annelida). Paratypes (N 5 2), one specimen fixed in formalin, dissected, same collection data as holotype (AMNH 5287, Annelida). One specimen fixed in 95% ethanol, Parc National Ranomafana, on trail between Talatakely and Vatoharanana (MG02-06), March 5, 2002, collected by Mark Siddall and Elizabeth Borda, (AMNH 5286, Annelida). DIAGNOSIS: Male pore in XI b5/b6; Female pore XII b5 (anterior); four complete annuli between gonopores; XXIV incomplete fourannulate; nine annuli between XXIII a2 and anus; oviducal glandular sac absent; muscular vaginal tube present. Prehensile lobe present. DESCRIPTION: Dorsum with dark brown to black background field, wide beige medial chain-link band, each link with darker field at center, approximately five annuli per link. Medial chain becomes continuous light field bordered with dark brown to black paramedial lines in posterior series. Paramedial dorsal light brown to white diamond-shaped patches at approximately the same position of every other chain-link of medial band. Dorsolateral light spots on sensillae of a2. Dark spots found along lateral margins, against light background that continues ventrally. Venter, light background with dark brown to black mottling. Annular papillae, present (figs. 8, 9). Duognathous, monostichodont jaws. Cephalic somites with five pairs eyespots, one on each of II, III, IV, V, and VI. Somites II, III, and IV uni-annulate, V and VI two-annulate and three-annulate, respectively, VII threeannulate. Midbody somites, VIII���XXIII, five-annulate. Nephropores, lateral, in furrow of b2/a2 of each somite, first nephropore observed at IX b2/a2. Posterior somites, XXIV incomplete four-annulate (b6 converges with b5 along the anterio-lateral margin of the respiratory auricle) and XXV���XXVII uni-annulate (fig. 10). Nine annuli (dorsally) between XXIII a2 and anus. Posteriolateral respiratory auricles, bilobed, and formed along the lateral margins of XXIV b5 to XXII (fig. 10). Caudal sucker, ventral, with 57 friction rays ventral on sucker. Prehensile lobe, present (fig. 11). General organization and regional morphology of male and female reproductive systems, not typical haemadipsoid (figs. 12��� 15). Male genital pore opens at the furrow of XI b5/b6 and female genital pore opens anteriorly in XII b5 (or a2/b5) (fig. 12). Gonopores separated by four complete annuli. Median reproductive system micromorphic (fig. 13). Median male reproductive system found entirely in XI (fig. 14). Male atrium, narrow, recurves anteriorly at XII. Short ejaculatory ducts exit the male atrium laterally in XI and give rise to small, globular ejaculatory bulbs in XI. Tightly coiled epididymes arise from ejaculatory bulbs, each twice the size of an ejaculatory bulb and recurved anteriad at XII. Median female reproductive system found entirely in XII (fig. 15). Single pair of globular ovaries, posteriorly directed in XII, with thin-walled oviducts that converge into common oviduct. Common oviduct is tightly coiled ventrally and inserts into a simple, muscular, vaginal tube. Vaginal tube recurves anteriad at approximately XIII, with tube widening anteriorly and terminating in bulb at the insertion point of the gonopore at XII b5. Haemadipsoid oviducal glandular sac absent. REMARKS: When Richardson (1975, 1978) included Haemadipsa dussumieri as a member of the genus Malagabdella, it was the only species that possessed four complete annuli between the gonopores. This was initially consistent with M. niarchosorum, n.sp. However, after the examination of the type specimen, it became obvious that M. dussumieri was distinct from M. niarchosorum, n.sp. (and from the genus Malagabdella). For example, the position of the female pore for the type specimen of H. dussumieri is on an obviously ventrally subdivided annulus of XII b5, whereas in M. niarchosorum, n.sp., it is not subdivided. In addition, as detailed above, M. dussumieri possesses characters that deviate from domanibdellid leeches. Therefore, with the exclusion of M. dussumieri from the Malagabdellinae, M. niarchosorum, n.sp. is distinct from all other members of the genus based on external coloration and patterns, the characteristics of the male and female reproductive systems, the position of the gonopores, and the posterior annulation series, but inclusive in the Malagabdellinae based on somital annulation pattern and the possession of 57 friction rays ventrally on the caudal sucker. ETYMOLOGY: This species is named after the Niarchos family in light of their generosity in funding this expedition. Without their support, the rediscovery of known species from Madagascar and the opportunity to describe another endemic species would not have been possible., Published as part of BORDA, ELIZABETH, 2006, A Revision of the Malagabdellinae (Arhynchobdellida: Domanibdellidae), with a Description of a New Species, Malagabdella niarchosorum, from Ranomafana National Park, Madagascar, pp. 1-16 in American Museum Novitates 3531 (1) on pages 7-9, DOI: 10.1206/0003-0082(2006)3531[1:AROTMA]2.0.CO;2, http://zenodo.org/record/4712713, {"references":["Richardson, R. L. 1975. A contribution to the general zoology of the land leeches (Hirudinea: Haemadipsoidea superfam. nov.). Acta Zoologica Academiae Scientiarum Hungaricae 21 (1 - 2): 119 - 152.","Richardson, R. L. 1978. On the zoological nature of land-leeches in the Seychelles Islands, and a consequential revision of the status of land-leeches in Madagascar (Hirudinea: Haemadipsoidea). Revue de Zoologie Africaine 92 (4): 837 - 866."]}

Malagabdella vagans (Blanchard, 1917) Figures 16���23 Haemadipsa vagans Blanchard, 1917: 665, fig. 14. Haemadipsa vagans ��� Soos, 1967: 426. Malagabdella vagans ��� Richardson, 1975: 142; Richardson, 1978: 860���863, fig. 4 A���D); Sawyer, 1986: 762. TYPE MATERIAL: Syntype, Madagascar ��� east coast forests, 1882, Lantz (MNHN 751; no. 143; N 5 1). Syntype, Madagascar ��� Montagne d���Ambre, Diego-Suarez, 1890, Dr. Catat? (MNHN 802; N 5 3). ADDITIONAL MATERIAL: Six specimens (AMNH 5288, Annelida) fixed in 95% ethanol and six specimens (AMNH 5289, Annelida) fixed in formalin (two specimens used for SEM), in forest adjacent to town of Andasibe,, 3 km from entrance of Analamazoatra (Perinet Special Reserve) (MG02-04), near the Antananarivo ��� Taomasina railroad, March 3, 2002. Brilliant green or orange when alive. Pale in color, almost neutral when fixed. Only markings found dorsally are paramedial wavy lines that become faded posteriorly. Dorsal and ventral papillae present (fig. 16). + male atrium (a), small bulbous ejaculatory bulbs (eb) and coiled epididymes (e). 15. Female reproductive system showing paired globular ovaries, each with independent oviducts that converge into the common oviduct (co) that inserts into a simple vaginal tube (vt). Duognathous, monostichodont jaws armed with serrated edge of minute teeth (fig. 17). Cephalic somites with five pairs of eyespots, one on each of II, III, IV, V, and VI. Somites II, III, and IV uni-annulate, V and VI twoannulate and three-annulate, respectively, VII three-annulate. Midbody somites VIII���XXIII five-annulate. Nephropores lateral in furrow of b2/a 2 in each somite, with first nephropore observed in IX b2/a2. XXIV��� XXVII uniannulate (fig. 18). Six annuli between XXIII a2 and anus. Respiratory auricles present, bilobed, and formed along the lateral margins of XXIV a2 to XXVII. Caudal sucker, ventral, with 58 friction rays ventral on sucker. Small prehensile lobe present (fig. 19). General organization and regional morphology of male and female reproductive systems not typical haemadipsoid (figs. 20���23). Male genital pore opens at the furrow of XI b5/b6 and the female genital pore opens at the furrow of XIII b1/b2 (fig. 20). Gonopores separated by seven complete annuli. Median reproductive system micromorphic and entirely in XI (figs. 21, 22). Male atrium bulbous. Short ejaculatory ducts exit the atrium ventrally in anterior XI and run laterally, posteriorly directed, giving rise to small ovoid ejaculatory bulbs. Epididymes, small, approximately the size of the ejaculatory bulbs, tightly coiled, and recurve anteriad at XII. Median female reproductive system micromorphic (fig. 23). Single pair of globular ovaries in XII, with short thin walled oviducts that coil and converge into the common oviduct. The common oviduct is tightly coiled in the median and exits in XIII. Oviducal glandular sac and muscular vaginal tube absent. REMARKS: Richardson (1975) noted a resemblance in the somital annulation pattern of H. vagans to Idiobdella seychellensis and, therefore, originally included H. vagans as a member of the genus Idiobdella. Richardson (1978) revised his previous classification of the leeches of Madagascar based on a specimen from the collection of the National Museum of Natural History that he identified as Malagabdella vagans (U.S. Nat. Mus. Cat. No. 55386; Acc. 242082; 2 km east of Perinet, Tamatave Province; off human skin). This resulted in the revision of the genus Malagabdella, the removal of the genus from Idiobdellidae, and the establishment of the Malagabdellinae (Richardson, 1978). Interestingly, Richardson (1978) reported that the specimen of M. vagans was engorged with blood and with the ������annulation obliterated������. Furthermore, it does not appear that he examined any type material for this species. Therefore, it is difficult to imagine that the annulation patterns he described and illustrat- ed for this species came from this specimen (see also M. fallax REMARKS), but rather a conversion of the annulation patterns described by Blanchard (1917). In his revision, Richardson (1978) described the male reproductive system for Malagasy leeches based on this USNM specimen of M. vagans. This resulted in an overgeneralization for the characteristics of male reproductive systems for the Malagabdella species. It should be mentioned that the observation of the male system was facilitated by the USNM specimen having been ������split through transversely at XII/ XIII providing access to the anterior portion of the male paired system and the male median region������ (Richardson, 1978: 863), yet he illustrates the majority of male system (i.e., atrium and ejaculatory bulbs and epididymes) found within XI., Published as part of BORDA, ELIZABETH, 2006, A Revision of the Malagabdellinae (Arhynchobdellida: Domanibdellidae), with a Description of a New Species, Malagabdella niarchosorum, from Ranomafana National Park, Madagascar, pp. 1-16 in American Museum Novitates 3531 (1) on pages 9-11, DOI: 10.1206/0003-0082(2006)3531[1:AROTMA]2.0.CO;2, http://zenodo.org/record/4712713, {"references":["Blanchard, R. 1917. Monographie des hemadipsines (sangsues terrestres). Bulletin de la Societe de Pathologie Exotique 10: 640 - 675.","Soos, A. 1967. Identification key to the leech (Hirudinoidea) genera of the world with a catalogue of species. IV. Family: Haemadipsidae. Acta Zoologica Academiae Scientiarum Hungaricae 13: 417 - 432.","Richardson, R. L. 1975. A contribution to the general zoology of the land leeches (Hirudinea: Haemadipsoidea superfam. nov.). Acta Zoologica Academiae Scientiarum Hungaricae 21 (1 - 2): 119 - 152.","Richardson, R. L. 1978. On the zoological nature of land-leeches in the Seychelles Islands, and a consequential revision of the status of land-leeches in Madagascar (Hirudinea: Haemadipsoidea). Revue de Zoologie Africaine 92 (4): 837 - 866.","Sawyer, R. T. 1986. Leech Biology and Behavior. Oxford: Clarendon Press."]}

Haemadipsa morsitans Blanchard, 1917: 664, fig. 13. Malagabdella morsitans ��� Richardson, 1975: 142; Richardson, 1978: 864, fig. 4E; Sawyer, 1986: 761. TYPE MATERIAL: Syntypes, Madagascar ��� Moramonga, east coast forests, 1893, Sikora? (MNHN 802; N 5 7). DESCRIPTION: Duognathous, monostichodont jaws. Cephalic somites with five pairs eyespots, one on each of II, III, IV, V, and VI. Somites II, III, and IV uni-annulate, V and VI two-annulate and three-annulate, respectively, VII three-annulate. Midbody somites VIII��� XXII five-annulate. Nephropores lateral in furrow of b2/a 2 in each somite. XXIII incomplete five-annulate (four annuli dorsally). XXIV incomplete three-annulate, XXV��� XXVII uni-annulate. Seven (eight) annuli between XXIII a2 and anus. Male genital pore at furrow of XI/XII and the female genital pore at furrow of XII/XIII. Gonopores separated by five complete annuli. REMARKS: Because I did not have access to fresh material of M. morsitans for dissection, the characteristics of the reproductive systems remain a mystery. Richardson���s (1978) redescription of this species was solely based on the somital annulation patterns and based on the position of the gonopores. The specimen Richardson (1978) examined (USNM 37970) was dried out and with a strongly compressed body. He also mentioned that the male pore was not obvious, but that the female pore was found in XII/XIII (i.e., ������two full annuli to a weakly defined end of the clitellum������; p. 864). The position of the gonopores has been confirmed based on the examination of the type specimen (see also M. fallax: REMARKS). Based on the latter and based on the annulation patterns renders M. morsitans distinct from other Malagabdella species, although the comparison of the reproductive systems to other species of Malagabdella is pending for a redescription of this species. Malagabdella niarchosorum, new species Figures 8���15 TYPE MATERIAL: Holotype, specimen fixed in 95% ethanol, Madagascar ��� Parc National Ranomafana in Talatakely (selectively logged, low montane forest; MG02-05), March 5, 2002. Collected by Mark Siddall, Elizabeth Borda, Evon Hekkala, Clara Imboule, and Raina Rakotondriany (AMNH 5285, Annelida). Paratypes (N 5 2), one specimen fixed in formalin, dissected, same collection data as holotype (AMNH 5287, Annelida). One specimen fixed in 95% ethanol, Parc National Ranomafana, on trail between Talatakely and Vatoharanana (MG02-06), March 5, 2002, collected by Mark Siddall and Elizabeth Borda, (AMNH 5286, Annelida). DIAGNOSIS: Male pore in XI b5/b6; Female pore XII b5 (anterior); four complete annuli between gonopores; XXIV incomplete fourannulate; nine annuli between XXIII a2 and anus; oviducal glandular sac absent; muscular vaginal tube present. Prehensile lobe present. DESCRIPTION: Dorsum with dark brown to black background field, wide beige medial chain-link band, each link with darker field at center, approximately five annuli per link. Medial chain becomes continuous light field bordered with dark brown to black paramedial lines in posterior series. Paramedial dorsal light brown to white diamond-shaped patches at approximately the same position of every other chain-link of medial band. Dorsolateral light spots on sensillae of a2. Dark spots found along lateral margins, against light background that continues ventrally. Venter, light background with dark brown to black mottling. Annular papillae, present (figs. 8, 9). Duognathous, monostichodont jaws. Cephalic somites with five pairs eyespots, one on each of II, III, IV, V, and VI. Somites II, III, and IV uni-annulate, V and VI two-annulate and three-annulate, respectively, VII threeannulate. Midbody somites, VIII���XXIII, five-annulate. Nephropores, lateral, in furrow of b2/a2 of each somite, first nephropore observed at IX b2/a2. Posterior somites, XXIV incomplete four-annulate (b6 converges with b5 along the anterio-lateral margin of the respiratory auricle) and XXV���XXVII uni-annulate (fig. 10). Nine annuli (dorsally) between XXIII a2 and anus. Posteriolateral respiratory auricles, bilobed, and formed along the lateral margins of XXIV b5 to XXII (fig. 10). Caudal sucker, ventral, with 57 friction rays ventral on sucker. Prehensile lobe, present (fig. 11). General organization and regional morphology of male and female reproductive systems, not typical haemadipsoid (figs. 12��� 15). Male genital pore opens at the furrow of XI b5/b6 and female genital pore opens anteriorly in XII b5 (or a2/b5) (fig. 12). Gonopores separated by four complete annuli. Median reproductive system micromorphic (fig. 13). Median male reproductive system found entirely in XI (fig. 14). Male atrium, narrow, recurves anteriorly at XII. Short ejaculatory ducts exit the male atrium laterally in XI and give rise to small, globular ejaculatory bulbs in XI. Tightly coiled epididymes arise from ejaculatory bulbs, each twice the size of an ejaculatory bulb and recurved anteriad at XII. Median female reproductive system found entirely in XII (fig. 15). Single pair of globular ovaries, posteriorly directed in XII, with thin-walled oviducts that converge into common oviduct. Common oviduct is tightly coiled ventrally and inserts into a simple, muscular, vaginal tube. Vaginal tube recurves anteriad at approximately XIII, with tube widening anteriorly and terminating in bulb at the insertion point of the gonopore at XII b5. Haemadipsoid oviducal glandular sac absent. REMARKS: When Richardson (1975, 1978) included Haemadipsa dussumieri as a member of the genus Malagabdella, it was the only species that possessed four complete annuli between the gonopores. This was initially consistent with M. niarchosorum, n.sp. However, after the examination of the type specimen, it became obvious that M. dussumieri was distinct from M. niarchosorum, n.sp. (and from the genus Malagabdella). For example, the position of the female pore for the type specimen of H. dussumieri is on an obviously ventrally subdivided annulus of XII b5, whereas in M. niarchosorum, n.sp., it is not subdivided. In addition, as detailed above, M. dussumieri possesses characters that deviate from domanibdellid leeches. Therefore, with the exclusion of M. dussumieri from the Malagabdellinae, M. niarchosorum, n.sp. is distinct from all other members of the genus based on external coloration and patterns, the characteristics of the male and female reproductive systems, the position of the gonopores, and the posterior annulation series, but inclusive in the Malagabdellinae based on somital annulation pattern and the possession of 57 friction rays ventrally on the caudal sucker. ETYMOLOGY: This species is named after the Niarchos family in light of their generosity in funding this expedition. Without their support, the rediscovery of known species from Madagascar and the opportunity to describe another endemic species would not have been possible. Malagabdella niarchosorum, new species Figures 8���15 TYPE MATERIAL: Holotype, specimen fixed in 95% ethanol, Madagascar ��� Parc National Ranomafana in Talatakely (selectively logged, low montane forest; MG02-05), March 5, 2002. Collected by Mark Siddall, Elizabeth Borda, Evon Hekkala, Clara Imboule, and Raina Rakotondriany (AMNH 5285, Annelida). Paratypes (N 5 2), one specimen fixed in formalin, dissected, same collection data as holotype (AMNH 5287, Annelida). One specimen fixed in 95% ethanol, Parc National Ranomafana, on trail between Talatakely and Vatoharanana (MG02-06), March 5, 2002, collected by Mark Siddall and Elizabeth Borda, (AMNH 5286, Annelida). DIAGNOSIS: Male pore in XI b5/b6; Female pore XII b5 (anterior); four complete annuli between gonopores; XXIV incomplete fourannulate; nine annuli between XXIII a2 and anus; oviducal glandular sac absent; muscular vaginal tube present. Prehensile lobe present. DESCRIPTION: Dorsum with dark brown to black background field, wide beige medial chain-link band, each link with darker field at center, approximately five annuli per link. Medial chain becomes continuous light field bordered with dark brown to black paramedial lines in posterior series. Paramedial dorsal light brown to white diamond-shaped patches at approximately the same position of every other chain-link of medial band. Dorsolateral light spots on sensillae of a2. Dark spots found along lateral margins, against light background that continues ventrally. Venter, light background with dark brown to black mottling. Annular papillae, present (figs. 8, 9). Duognathous, monostichodont jaws. Cephalic somites with five pairs eyespots, one on each of II, III, IV, V, and VI. Somites II, III, and IV uni-annulate, V and VI two-annulate and three-annulate, respectively, VII threeannulate. Midbody somites, VIII���XXIII, five-annulate. Nephropores, lateral, in furrow of b2/a2 of each somite, first nephropore observed at IX b2/a2. Posterior somites, XXIV incomplete four-annulate (b6 converges with b5 along the anterio-lateral margin of the respiratory auricle) and XXV���XXVII uni-annulate (fig. 10). Nine annuli (dorsally) between XXIII a2 and anus. Posteriolateral respiratory auricles, bilobed, and formed along the lateral margins of XXIV b5 to XXII (fig. 10). Caudal sucker, ventral, with 57 friction rays ventral on sucker. Prehensile lobe, present (fig. 11). General organization and regional morphology of male and female reproductive systems, not typical haemadipsoid (figs. 12��� 15). Male genital pore opens at the furrow of XI b5/b6 and female genital pore opens anteriorly in XII b5 (or a2/b5) (fig. 12). Gonopores separated by four complete annuli. Median reproductive system micromorphic (fig. 13). Median male reproductive system found entirely in XI (fig. 14). Male atrium, narrow, recurves anteriorly at XII. Short ejaculatory ducts exit the male atrium laterally in XI and give rise to small, globular ejaculatory bulbs in XI. Tightly coiled epididymes arise from ejaculatory bulbs, each twice the size of an ejaculatory bulb and recurved anteriad at XII. Median female reproductive system found entirely in XII (fig. 15). Single pair of globular ovaries, posteriorly directed in XII, with thin-walled oviducts that converge into common oviduct. Common oviduct is tightly coiled ventrally and inserts into a simple, muscular, vaginal tube. Vaginal tube recurves anteriad at approximately XIII, with tube widening anteriorly and terminating in bulb at the insertion point of the gonopore at XII b5. Haemadipsoid oviducal glandular sac absent. REMARKS: When Richardson (1975, 1978) included Haemadipsa dussumieri as a member of the genus Malagabdella, it was the only species that possessed four complete annuli between the gonopores. This was initially consistent with M. niarchosorum, n.sp. However, after the examination of the type specimen, it became obvious that M. dussumieri was distinct from M. niarchosorum, n.sp. (and from the genus Malagabdella). For example, the position of the female pore for the type specimen of H. dussumieri is on an obviously ventrally subdivided annulus of XII b5, whereas in M. niarchosorum, n.sp., it is not subdivided. In addition, as detailed above, M. dussumieri possesses characters that deviate from domanibdellid leeches. Therefore, with the exclusion of M. dussumieri from the Malagabdellinae, M. niarchosorum, n.sp. is distinct from all other members of the genus based on external coloration and patterns, the characteristics of the male and female reproductive systems, the position of the gonopores, and the posterior annulation series, but inclusive in the Malagabdellinae based on somital annulation pattern and the possession of 57 friction rays ventrally on the caudal sucker. ETYMOLOGY: This species is named after the Niarchos family in light of their generosity in funding this expedition. Without their support, the rediscovery of known species from Madagascar and the opportunity to describe another endemic species would not have been possible., Published as part of BORDA, ELIZABETH, 2006, A Revision of the Malagabdellinae (Arhynchobdellida: Domanibdellidae), with a Description of a New Species, Malagabdella niarchosorum, from Ranomafana National Park, Madagascar, pp. 1-16 in American Museum Novitates 3531 (1) on pages 6-9, DOI: 10.1206/0003-0082(2006)3531[1:AROTMA]2.0.CO;2, http://zenodo.org/record/4712713, {"references":["Blanchard, R. 1917. Monographie des hemadipsines (sangsues terrestres). Bulletin de la Societe de Pathologie Exotique 10: 640 - 675.","Richardson, R. L. 1975. A contribution to the general zoology of the land leeches (Hirudinea: Haemadipsoidea superfam. nov.). Acta Zoologica Academiae Scientiarum Hungaricae 21 (1 - 2): 119 - 152.","Richardson, R. L. 1978. On the zoological nature of land-leeches in the Seychelles Islands, and a consequential revision of the status of land-leeches in Madagascar (Hirudinea: Haemadipsoidea). Revue de Zoologie Africaine 92 (4): 837 - 866.","Sawyer, R. T. 1986. Leech Biology and Behavior. Oxford: Clarendon Press."]}