Field of Science

The Mouse Shrews of Africa

Shrews are one of the less appreciated groups of mammals. Small (some are among the smallest mammals on earth), skulking, they are often overlooked but are nevertheless represented by a diversity of species in many parts of the world. Among this diversity are the mouse shrews of the genus Myosorex.

Forest shrew Myosorex varius, from Roberts (1951).


Myosorex is a genus of nearly twenty known species of shrew, some of which have only been identified very recently, with more probably yet to be described. They are known in the modern fauna only from sub-Saharan Africa, though the fossil record indicates they once extended as far north as Spain (Furió et al. 2007). Mouse shrews differ from most other living shrew genera (except the closely related Congosorex) in the presence of a tiny vestigial tooth in the lower jaw behind the first antemolar (the tooth behind the incisors, so called in shrews because it is unclear whether it corresponds to a canine or premolar in relation to the teeth of other mammals). Because their teeth lack the red pigment found in shrews of the subfamily Soricinae, Myosorex have historically been classified with the white-tooth shrews of the Crocidurinae. However, the presence of white teeth is, of course, a primitive feature of questionable significance phylogenetically. Instead, more recent authors have pointed to the retention of the second antemolar and other features to support recognising Myosorex and related African shrew genera as relictual members of a third subfamily, the Myosoricinae, that may also include a number of earlier fossil shrews (this group has also been known as the 'Crocidosoricinae' but 'Myosoricinae' is the name with priority; the argument by Furió et al., 2007, that the latter name cannot be used for the broader subfamily because it was originally used only for the African genera has no standing under current nomenclatorial rules).

Foraging forest shrew, copyright Johnny Wilson.


In the modern fauna, Myosorex species have a very scattered distribution. Species found in central and eastern Africa are restricted to high mountains, among moist, densely vegetated environments. Species found in southern Africa are often found in similar habitats. However, the species M. varius is also found in drier locations at lower altitudes, closer to the South African coast. Nevertheless, it is still restricted to areas with high seasonal rainfall, or (in part of Western Cape Province) zones dominated by succulent vegetation where low levels of actual rainfall may be compensated for by precipitation from mist (Meester 1958). The distribution of the genus as a whole is marked by a broad gap of over 1600 km separating the northern limit of species in South Africa and Zimbabwe from their nearest neighbours to the north in the DRC and Kenya, a gap that they were presumably only able to cross in the past when climate conditions were more amenable.

This sensitivity to environment means that Myosorex species may be very vulnerable to changes in habitat. Several species are restricted to limited ranges and several are recognised as potentially endangered. The prospect of climate change makes this vulnerability even worse: as levels of rainfall decrease, mouse shrews will be forced to retreat to ever higher altitudes, and there's only so high they can go before running out of mountain.

REFERENCES

Furió, M., A. Santos-Cubedo, R. Minwer-Barakat & J. Agustí. 2007. Evolutionary history of the African soricid Myosorex (Insectivora, Mammalia) out of Africa. Journal of Vertebrate Paleontology 27 (4): 1018–1032.

Meester, J. 1958. Variation in the shrew genus Myosorex in southern Africa. Journal of Mammalogy 39 (3): 325–339.

Five-fingers and Lancewoods

Longtime readers of this blog will know that my knowledge of plants has always been fairly rudimentary. As a young'un, I only ever learnt to distinguish some of the more common and visible varieties. As a student, I did take a few botany classes, but only really enough to learn that plant biology is complicated and terrifying. Since then, I've continued in much the same vein. But for today's post, I'm looking at something I do recall being aware of in my youth: the lancewoods and five-fingers of the genus Pseudopanax.

Horticultural variant of coastal five-finger Pseudopanax lessonii, copyright Leonora Enking.


Pseudopanax is a genus of a dozen species of small tree (mostly growing about five to seven metres in height) found only in New Zealand (Perrie & Shepherd 2009). Various species have also been assigned to the genus from locations around the Pacific (China, Tasmania, New Caledonia and Chile) but recent studies have lead to their exclusion. A handful of New Zealand species previously included in Pseudopanax have also been separated as the genus Raukaua (Mitchell et al. 1997). The historical taxonomy of the group is confusing, with species being variously attributed to genera Panax, Nothopanax, Neopanax and Polyscias. Things seem to have settled down a bit in recent years but there is still the possibility we may one day see Neopanax rise again (Perrie & Shepherd 2009).

Chatham Islands lancewood Pseudopanax chathamicus, copyright Krzysztof Ziarnek, Kenraiz.


Pseudopanax belongs to the family Araliaceae, a group that is primarily composed of tropical and subtropical shrubs and trees. Araliaceae are commonly referred to as "the ivy family", after one of their best-known members, the common ivy Hedera helix, but, as is not uncommon when a tropical family gets named after one of their European outliers, ivy is pretty weird by Araliaceae standards. Pseudopanax species are perhaps a bit more typical. They have large leaves, often more or less toothed or lobed along the margins. In a number of species, the leaves are palmately divided into three or five separate leaflets, hence the aforementioned vernacular name of 'five-finger'. In one group of species, the lancewoods, the lateral leaflets have been lost and the now undivided leaf is more or less long and narrow. Hybrids between five-fingers and lancewoods may have multiple leaflets like a five-finger but the leaflets shaped like those of a lancewood; New Zealand botanist Leon Perrie has written a post about hybridisation in this genus that you can read here. The trees are usually dioecious (male and female flowers are borne on separate trees) and the individually small flowers are borne aggregated in compound umbels. Fruits are fleshy berries.

Collection of lancewoods P. crassifolius showing the variation in leaf form, copyright Petra Gloyn. Two individuals on the left are young tress with hanging leaves; to the right is a more mature individual with spreading leaves.


Within Pseudopanax, the lancewoods are particularly renowned for their exhibition of heteroblasty, a phenomenon where the appearance of the leaves changes significantly as the tree matures. Juvenile leaves of the common lancewood P. crassifolius and toothed lancewood P. ferox are remarkably long, slender, strongly toothed along the margin, stiff and leathery, and hang downwards around the young tree like a skirt. As the tree approaches its mature height, it starts producing shorter, softer, less serrate leaves that are held in a more or less horizontal position.

Changes in growth habit with maturity seem to be surprisingly common among New Zealand plants and there has been a lot of discussion about why this might be. One suggestion that has certainly received a lot of public attention is that it is a relic of browsing by the large herbivorous birds such as moa that dominated the New Zealand environment prior to human settlement. Juvenile plants developed a habit that was energetically expensive but discouraged browsing by birds; as they grew high enough to escape the reach of such browsers, they changed to a less demanding form. I personally tend to be skeptical of these kinds of claims of historical baggage, not least because the extinction of one-half of the equation makes them very hard to test in any way, but I will admit that this case does perhaps have a bit more credibility than, for instance, claims elsewhere of giant fruits being dependent on long-extinct megafauna. Alternatively, it has been suggested that changes in growth habit may be related to climatic conditions; the juvenile leaves of P. crassifolius dissipate heat more effectively than those of mature trees (Gould 1993). Heteroblasty is less pronounced in the montane lancewood P. linearis of the South Island and almost absent in the Chatham Islands lancewood P. chathamicus, an insular derivative of P. crassifolius. Were these species insulated from the selective pressures affecting the other two? It should also be pointed out that the two proposals mentioned here are not mutually exclusive; the consideration of one as a factor does not automatically rule out the other.

REFERENCES

Gould, K. S. 1993. Leaf heteroblasty in Pseudopanax crassifolius: functional significance of leaf morphology and anatomy. Annals of Botany 71: 61–70.

Mitchell, A. D., D. G. Frodin & M. J. Heads. 1997. Reinstatement of Raukaua, a genus of the Araliaceae centred in New Zealand. New Zealand Journal of Botany 35 (3): 309–315.

Perrie, L. R., & L. D. Shepherd. 2009. Reconstructing the species phylogeny of Pseudopanax (Araliaceae), a genus of hybridising trees. Molecular Phylogenetics and Evolution 52: 774–783.

Chilostomellidae: Deep Forams

Holotype of Chilostomella serrata, from the Smithsonian National Museum of Natural History.


The specimen in the figure above is a fairly typical representative of the Chilostomellidae, a cosmopolitan family of forams known from the Jurassic to the present day. Members of this family have a translucent calcareous test with chambers arranged in a trochospiral (broad conical) or planispiral (flat spiral) pattern. The chambers of each spiral are expanded to cover over the prior spirals so only the outermost spiral is generally visible. The aperture of the test in the final chamber is a narrow slit along the margin with the underlying chamber (Loeblich & Tappan 1964).

Despite their long history and wide distribution, I get the general impression that chilostomellids are not usually abundant. They are generally restricted to deeper waters, more than 100 m below the surface (Cushman et al. 1954). Members of the genus Chilostomella, at least, have commonly been regarded as associated with low-oxygen environments. However, it has also been suggested that their favoured conditions are not so much a question of low oxygen as high organic flux (Jorissen 2002). Perhaps the best location to find chilostomellids would be around sites where dead animals and seaweeds have fallen to the deeper waters below.

REFERENCES

Cushman, J. A., R. Todd & R. J. Post. 1954. Recent Foraminifera of the Marshall Islands. Bikini and nearby atolls, part 2, oceanography (biologic). Geological Survey Professional Paper 260-H: 319–384, pls 82–93.

Jorissen, F. J. 2002. Benthic foraminiferal microhabitats below the sediment-water interface. In: Sen Gupta, B. K. (ed.) Modern Foraminifera pp. 161–179. Kluwer Academic Publishers: Dordrecht.

Loeblich, A. R., Jr & H. Tappan. 1964. Treatise on Invertebrate Paleontology pt C. Protista 2. Sarcodina: chiefly "thecamoebians" and Foraminiferida vol. 2. The Geological Society of America, and The University of Kansas Press.

Digging for Tellina

When I was a child, a large part of my extended family would gather over the Christmas period to park their tents and caravans alongside the estuary downhill from my great-grandparents' house (in the usual way of these things, my memory has these summer camping periods lasting for ages, but I don't think they could have been longer than a week or so). While we were there, I would spend a fair chunk of the day looking for the wildlife that inhabited the slightly muddy estuary beach. Among these were various bivalves whose shells could be found littering the shoreline, or which might be found by digging in the sand at low tide. Close to the surface were New Zealand cockles Austrovenus stutchburyi (not actually a direct relative of the English cockle but a member of the Veneridae family that has adopted a similar body form). A little deeper were pipis and tuatuas. And a little deeper again were the flat, slender shells of Tellina.

Thin tellin Tellina tenuis, copyright S. Rae.


I should note that Tellina species are not really deep burrowers in the grand scheme of things, generally only embedding themselves about one to three centimetres below the surface, but again I must ask that you make allowances for childhood memories. Their low profile and weakly inflated shells also make them fast diggers so they were probably able to elude most casual explorations. Like most subsurface bivalves, Tellina species are sediment feeders. Their usual aspect is lying horizontally beneath the sediment, extending their long, unfused siphons to the surface to gather detritus (Ujino & Matuskuma 2010; the shells of Tellina are usually twisted slightly to one side at the end to facilitate the siphons' passage). Even if you've not seen the Tellina animals themselves, you may have seen the radiating trails made by the siphons as they extend along the top of the sediment.

Sunrise tellin Tellina radiata, copyright James St. John.


Tellina is an extremely diverse genus, with species found worldwide and recognised through the entirety of the Mesozoic (Moore 1969). These species vary greatly in appearance, with shells varying from almost completely smooth to strongly ornamented, and from subcircular to quite elongate. It should therefore come as little surprise that numerous attempts have been made to divide Tellina between various subgenera and genera but issues such as homoeomorphy in Tellina's evolution (where distinct lineages have converged on similar body forms) have lead to disagreement over the best system to adopt. In 1934, the malacologist A. E. Salisbury complained that, "The number of genera, subgenera, and sections into which the Tellinidae has been cut up is getting somewhat appalling; the list of names is still increasing every year, and, if every variation of form is magnified, it is quite possible to go on until at last each species becomes the representative of a different genus and each variety that of a subgenus" (of course, as seems almost inevitable when one encounters complaints of this kind, Salisbury himself then proceeds to add to the tally of generic names in that same paper). Though I suspect most modern malacologists would probably disagree with the extremely broad concept of Tellina advocated by Salisbury, the question of how best to handle the genus taxonomically remains an open one.

REFERENCES

Moore, R. C. (ed.) 1969. Treatise on Invertebrate Paleontology pt N. Mollusca 6. Bivalvia vol. 2. The Geological Society of America, Inc., and The University of Kansas.

Salisbury, A. E. 1934. On the nomenclature of Tellinidae, with descriptions of new species and some remarks on distribution. Proceedings of the Malacological Society of London 21: 74–91.

Ujino, S., & A. Matsukuma. 2010. Inverse life positions of three species in the genus Cadella (Bivalvia: Tellinidae). Molluscan Research 30 (1): 25–28.

Agenioideus: Average Spider Hawks

I have commented in earlier posts on the challenges of identifying spider hawks of the family Pompilidae, resulting from this wasp family's combination of high species diversity with a mostly conservative body plan. As a result of this conservatism, pompilid classification has tended to drift towards a situation where the majority of species are included in a relatively small number of somewhat vaguely defined genera. Each of the species included in one of these genera can be associated with other species in the genus, and groups of species approach each other closely enough that clear lines cannot be settled upon, but identifying features shared by all members of the genus can prove difficult. A good example of one such genus is Agenioideus.

Female Agenioideus birkmanni, from the University of Texas at Austin.


Species assigned to Agenioideus can be found pretty much worldwide though the greatest diversity occurs in warmer parts of the Holarctic. Though there does not seem to be a great deal of disagreement over which species should be placed in this genus, it seems a little difficult to say exactly what makes an Agenioideus. If anything, Agenioideus species seem to be associated by how relentlessly average they are, possessing a unique combination of characters that are none of them individually unique. They have wings with three submarginal cells, a broad metapostnotum in front of the propodeum, and legs ending in a small arolium with a weak comb of setae between a pair of long claws, mostly with a single small ventral tooth (Krogmann & Austin 2012). If you don't know exactly what those terms mean, just know that they are all quite unspecialised features for pompilids. Males often have asymmetrical claws on the forelegs, with the inner claw strongly bent and bifid while the outer claw is like those on the other legs, and the pterostigma (the dark node at the front of the fore wings) is relatively large compared to other genera. Females often have a comb of longer spines on the inner margin of the fore tarsi. But these last, more derived, features may not be universally present across all species of the genus.

Female Agenioideus nigricornis with redback spider Latrodectus hasselti as prey, copyright Mark Newton.


As befits their unspecialised appearance, most Agenioideus species (as far as we know) are relatively unspecialised in their nesting behaviour (Shimizu 1997). Like other pompilids, they lay their eggs on paralysed spiders that will provide food for the larva when it hatches. Most Agenioideus species construct simple nests with a single brood cell containing a single spider for each nest. One European species, A. nubecula, is known to produce slightly more extensive nests with up to four cells. The nest may be made by digging in loose soil or by using a pre-existing cavity; whether the wasp is more likely to do one or the other is correlated with whether she possesses a well-developed tarsal comb. A Japanese species, A. ishikawai, is known to at least partially dig a nest before capturing a spider, completing construction after bringing it back. The most specialised provisioning behaviour known for the genus, however, is found in another European species, A. coronatus. This species hunts jumping spiders which she paralyses with her sting as is standard. The paralysis, however, is only temporary, lasting just a few minutes, just long enough for the female to deposit an egg near the base of the spider's abdomen where it cannot easily remove it. The spider is then freed to go about its business without being placed in a nest, until the wasp larva hatches and feeds on its host in the manner of a parasitoid.

REFERENCES

Krogmann, L., & A. D. Austin. 2012. Systematics of Australian Agenioideus Ashmead (Hymenoptera: Pompilidae) with the first record of a spider wasp parasitizing Latrodectus hasselti Thorell (redback spider). Australian Journal of Entomology 51: 166–174.

Shimizu, A. 1997. Taxonomic studies on the Pompilidae occurring in Japan north of the Ryukyus: the genus Agenioideus Ashmead (Hymenoptera). Japanese Journal of Entomology 65 (1): 143–167.

Morion Revisited

In an earlier post, I introduced you to the carabid beetle genus Morion, currently recognised as including about forty species from tropical and subtropical regions around the world. In that post, I mentioned how Will (2003) had questioned the monophyly of this genus, owing to its lack of derived features in comparison with closely related genera. In this post, I'll take the opportunity to dive a little further into ways the genus may be divided.

Just to remind you what we're looking at: Morion monilicornis, copyright Robert Webster.


As noted by Will (2003), many authors have recognised two subgenera within Morion, Morion sensu stricto and Neomorion. These subgenera were first established by Jeannel (1948) who identified a distinction between species he examined from the Old World (Africa and Asia) and the Americas. The Old World species, to which Jeannel gave the name Neomorion, had a number of setae along the rear margin of the last ventrite of the abdomen. In males, the basal segment of the fore tarsus had the inner apex drawn out into a tooth. The aedeagus bears a large dorsoapical orifice; in Old World Morion examined by Jeannel, this orifice was covered over by a large bilobed lamella. In the New World species of Morion sensu stricto, in contrast, the rear margin of the last ventrite bore only two setae, and males lacked a medioapical tooth on the basal fore tarsomere. The opening of the aedeagus lacked a covering lamella. In his key to Morion and related genera, Will (2003) referred only to the state of the male fore tarsus as distinguishing the subgenera. It may be that this indicates that the significance of the other characters described by Jeannel had been subject to question, but I suspect that they may have omitted by Will because their state in a number of Morion species remains unknown.

Aedeagi of Morion in lateral and dorsal view from Jeannel (1948). On the left is an Old World species (Morion orientale), on the right a South American species (M. georgiae).


A particular notable lacuna in Jeannel's brief survey of Morion was that he didn't look at any Australian species. In his description of M. crassipes*, a species found in the vicinity of Cairns in Queensland, Sloane (1904) noted that the male fore tarsi differed from those of other Australian species in having the "basal joints rounded and not produced at inner apical angle", implying that most Australian Morion have tarsi resembling those of the New World species. Moore (1965), in a review of Australian genera of Pterostichinae, describes Morion as having an aedeagus with an orifice on the dorsum, with no mention of a covering lamella, again also suggesting a resemblance to New World rather than Old World species (unfortunately, Moore did not specify exactly which species his description of the genitalia was based on). Moore (1965) also noted that Australian species were distinctive among Morion in having a pronotum with more than the two setae along each lateral margin found in species from elsewhere. One species which is found in New Guinea and northern Australia, M. longipennis, does have only two pairs of marginal setae on the pronotum, and Darlington (1962) suggested that it was probably more closely related to Asian species than to other Australian Morion. The aforementioned M. crassipes differs from other Australian species in a number of significant features, including large size (it grows to a full inch in length) and modified legs, and Sloane (1904) did briefly wonder whether it should even be regarded as a Morion, but it does share the plurisetose pronotal margins. I should note that I've found no reference to the pilosity of the last ventrite in any Australian species.

*Under the name 'Morio crassipes'; confusion about whether this genus should be called Morion or Morio lingered for a long time in the early 20th century.

So overall, there is the suggestion of three distinct groups within Morion, for the Old World, American, and Australian species, with the last two more similar to each other than to the first. The New World species of Morion are more diverse in South America than in North America, and one might be tempted to line up the relationship between the three groups with the division of Gondwana. The Old World species group may have diverged first with the separation of Africa and/or India, followed by the South American and Australian lineages diverging as their own continents became isolated. The South American species group may have spread into North America with the formation of the Central American land bridge, and the increased proximity of Australasia to Asia may have allowed members of the Old World group such as M. longipennis to invade from the northwest. However, I've based this scenario on some pretty weak assumptions based on very incomplete data, and it would require a more detailed investigation before we could say if there's any merit to it.

REFERENCES

Darlington, P. J., Jr. 1962. The carabid beetles of New Guinea. Part I. Cicindelinae, Carabinae, Harpalinae through Pterostichini. Bulletin of the Museum of Comparative Zoology 126 (3): 319–564, 4 pls.

Jeannel, R. 1948. Faune de l'Empire Français. X. Coléoptères Carabiques de la Région Malgache (deuxième partie). Office de la Recherche Scientifique Coloniale: Paris.

Moore, B. P. 1965. Studies on Australian Carabidae (Coleoptera). 4.—The Pterostichinae. Transactions of the Royal Entomological Society of London 117 (1): 1–32.

Sloane, T. G. 1904. Studies in Australian entomology. No. XIV. New species of geodephagous Coleoptera from tropical Australia. Cicindelidae (3), and Carabidae (5) [Platysmatini, Morioni, Perigonini, Masoreini, and Physocrotaphini]. Proceedings of the Linnean Society of New South Wales 29 (3): 527–538.

Will, K. W. 2003. Review and cladistic analysis of the generic-level taxa of Morionini Brullé (Coleoptera: Carabidae). Pan-Pacific Entomologist 79 (3–4): 212–229.

Pseudogagrella: A Harvestman Torn

The Sclerosomatidae are one of the most diverse of the currently recognised harvestmen families, and one of the most problematic when it comes to classification. In various earlier posts, I have noted the challenges that bedevil sclerosomatid systematics, many reflecting a historical focus on superficial external features of questionable evolutionary significance. Perhaps no taxon more neatly exemplifies the problems with higher sclerosomatid classification than the eastern Asian genus Pseudogagrella.

Pseudogagrella sakishimensis, copyright Tomoya Suzuki.


Historically, the greater number of sclerosomatids have been divided between two major subfamilies, the Leiobuninae and Gagrellinae. The Leiobuninae have mostly been recognised as living in the northern temperate regions whereas the Gagrellinae were mostly tropical. The division between the two subfamilies has long been regarded as more than a little fuzzy, and has usually hinged on a single feature: the presence (Gagrellinae) or absence (Leiobuninae) of rings of flexible integument (noduli) in the femora of the legs. Pseudogagrella is a genus of sclerosomatid harvestmen recognised from Japan, Taiwan, China and Sumatra (Chen & Shih 2017). Members of this genus lack leg noduli so have historically been included in the Leiobuninae. The problem is that their overall appearance, with a tendency to bold coloration, a tall median spine rising from the hardened scute covering most of the abdomen, and legs that are not merely long but ludicrously so (even by harvestman standards), is extremely similar to species of Gagrellinae. So much so, in fact, that some species currently placed in Pseudogagrella were long included in the archetypical gagrelline genus, Gagrella (Suzuki 1977).

With the distinction between the two subfamilies being so vague, I don't think it really came as that much surprise to anyone when molecular phylogenetics underlined the need for a thorough re-working of sclerosomatid systematics. Though the analysis conducted by Hedin et al. (2012) did not support the prior distinction between 'leiobunines' and 'gagrellines', it did suggest the existence of distinct lineages occupying distinct geographical regions. One species of Pseudogagrella included in the analysis (the southern Japanese P. amamiana) was placed in a cluster of eastern Asian species including other Asian 'gagrellines', but also the 'leiobunine' 'Leiobunum' japonicum. So it seems likely that, should subfamilies of Sclerosomatidae continue to be recognised, Pseudogagrella will indeed be a member of Gagrellinae, but Gagrellinae itself shall not quite be what people think of it as being.

Pseudogagrella dorsomaculata, copyright Tyus Ma.


There is also, of course, the question of whether Pseudogagrella itself is a coherent unit. Hedin et al. (2012) included only the one Pseudogagrella species in their analysis and the need for an extensive revision of the Asian sclerosomatid genera still remains. A study of Chinese species assigned to the genus Melanopa (Zhang & Zhang 2013), which is primarily distinguished from Gagrella by having relatively shorter legs, suggested the possibility of this 'genus' being divided between groups of Palaearctic and Indo-Malayan species, and I've wondered if this division might carry further (unfortunately, I'm not aware of any Indo-Malayan 'gagrellines' being included in molecular phylogenies; I think all the Asian species covered by Hedin et al. were Palaearctic). The majority of Pseudogagrella species, found in Japan and Taiwan, can be comfortably compared to other sclerosomatids from that region, but the Sumatran P. multimaculata, and possibly the southern Chinese species, might turn out to be closer to their own geographical peers. As always, a great deal of research remains to be done.

REFERENCES

Chen, S.-L., & H.-T. Shih. 2017. Descriptions of three new species of the harvestmen genus Pseudogagrella (Opiliones: Sclerosomatidae: Gagrellinae) from Taiwan, supported by morphological and molecular evidence. Zootaxa 4268 (1): 34–52.

Hedin, M., N. Tsurusaki, R. Macías-Ordóñez & J. W. Shultz. 2012. Molecular systematics of sclerosomatid harvestmen (Opiliones, Phalangioidea, Sclerosomatidae): geography is better than taxonomy in predicting phylogeny. Molecular Phylogenetics and Evolution 62 (1): 224–236.

Suzuki, S. 1977. Opiliones from Taiwan (Arachnida). Journal of Science of the Hiroshima University, Series B, Division 1 (Zoology) 27 (1): 121–157.

Zhang, C., & F. Zhang. 2013. Notes on some species of the genus Melanopa (Opiliones: Sclerosomatidae: Gagrellinae) from China, with description of a new species. Journal of Arachnology 41: 306–318.