The Order Gryllioblattodea Contains a Single Family, the ____________.
PLoS 1. 2010; 5(ix): e12850.
A Second New Species of Ice Crawlers from China (Insecta: Grylloblattodea), with Thorax Development and the Prediction of Potential Distribution
Ming Bai
1 Key Laboratory of Zoological Systematics and Development, Constitute of Zoology, Chinese University of Sciences, Beijing, People'south Republic of Prc,
Karl Jarvis
ii Northern Arizona Academy, Flagstaff, Arizona, U.s. of America,
Shu-Yong Wang
ane Key Laboratory of Zoological Systematics and Development, Institute of Zoology, Chinese Academy of Sciences, Beijing, People's Democracy of China,
Ke-Qing Song
ane Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Beijing, Cathay,
Yan-Ping Wang
ane Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese University of Sciences, Beijing, People'southward Commonwealth of China,
Zhi-Liang Wang
1 Fundamental Laboratory of Zoological Systematics and Evolution, Plant of Zoology, Chinese University of Sciences, Beijing, People's Republic of Communist china,
Wen-Zhu Li
1 Fundamental Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Beijing, People's Democracy of Mainland china,
Wei Wang
3 Foreign Economical Cooperation Part, Ministry of Environmental Protection, Beijing, People's Commonwealth of China,
Xing-Ke Yang
1 Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese University of Sciences, Beijing, People's Democracy of Prc,
Robert DeSalle, Editor
Received 2010 Apr 15; Accepted 2010 Jul 8.
Abstract
Modern grylloblattids are one of the least diverse of the modern insect orders. The thorax changes in morphology might exist associated with the changes of the function of the forelegs, wing loss, changes in beliefs and adaptation to habitat. As temperature is the main barrier for migration of mod grylloblattids, the range of each species is extremely limited. The potential distribution areas of grylloblattids remain unclear. A second new species of ice crawlers (Insecta: Grylloblattodea), Grylloblattella cheni Bai, Wang et Yang sp. nov., is described from China. The distribution map and key to species of Grylloblattella are given. A comparison of the thorax of extant and extinct Grylloblattodea is presented, with an emphasis on the pronotum using geometric morphometric analysis, which may reverberate thorax adaptation and the development of Grylloblattodea. Potential global distribution of grylloblattids is inferred. Highly diversified pronota of extinct Grylloblattodea may reflect diverse habitats and niches. The relatively homogeneous pronota of modern grylloblattids might be explained by ii hypotheses: synapomorphy or convergent evolution. Most fossils of Grylloblattodea contain an patently longer meso- and metathorax than prothorax. The length of the meso- and metathorax of modernistic grylloblattids is unremarkably shorter than the prothorax. This may be associated with the wing loss, which is accompanied by musculus reduction and changes to the thoracic skeleton system. Threats to grylloblattids and several conservation comments are too provided.
Introduction
Modern grylloblattids (too known as ice bugs, water ice crawlers, and rock crawlers), all occur north of ∼35° latitude in cool-temperate areas of the U.s., Canada, Russia, Japan, Korea and Prc, and they are restricted to cold and extreme habitats that are difficult to admission. They are ane of the least diverse of the modernistic insect orders, consisting of 29 species, including a new species described below. All of the known extant species, which belong to the family unit Grylloblattidae and 5 genera, Galloisiana, Grylloblattina, Grylloblattella, Namkungia and Grylloblatta. Water ice crawlers tin be considered as "living fossils" with presently relict distributions [one], [ii]. Grylloblattids are generally found on northward-facing talus slopes, snowfall patches about forest at high elevations (1500–3000 m), in caves with permanent water ice at low elevations (300–1000 m) [three], [four], and some Grylloblattina are from 5 grand–300 g, much lower than near other grylloblattids [five]. They live on and in soil, in caves, and beneath stones and in crevices of mountainous regions. They are principally feces feeders on other insects, though they will consume plant material, mucus, and detritus [half-dozen].
Grylloblattids are extremely rare in China. Mr. Shu-Yong Wang collected the first grylloblattid from China, which was ane male Galloisiana sinensis Wang, 1987 [7] specimen from Mt. Changbaishan, Jilin in 1986. Over 20 years afterward, Mr. Ke-Qing Song collected the second grylloblattid from China, which is one female Grylloblattella cheni sp. november. in Akekule Lake (White Lake), Xinjiang, China. The inclusion of this species to Grylloblattella expands the genus to 3 species, the other two species existence constitute in western to cardinal Siberia, Russia: K. pravdini in the Altai Mountains and G. sayanensis in the Sayan Mountains (Fig. 1C).
Grylloblattella cheni Bai, Wang et Yang sp. nov.
(A) Female. (B) Habitat. (C) Type localities of all known three species of Grylloblattella.
Modernistic grylloblattids are xiv–34 mm long, wingless, pale, and either nocturnal or cavernicolous. Adults have long cerci with five–ten segments, and females have a sword-shaped ovipositor similar in shape to that of katydids (Orthoptera: Tettigoniidae). The single extant family can exist contrasted with 46 families described from the fossil record, which extend to the Late Carboniferous [8], [9], [10]. The morphology of grylloblattodeans was stable with merely pocket-size changes during the ∼300 Million years of development, except thorax variations, which are the virtually significant difference between extant and extinct members of Grylloblattodea. The thorax, which contains the muscles of the legs and wings, had inverse in some caste during the development of Grylloblattodea. This might be associated with the changes of the function of the forelegs, fly loss, changes in behavior and adaptation to habitat. Here we present a comparison of the thorax of extant and extinct Grylloblattodea, with an emphasis on the pronotum using geometric morphometric analysis, which may reverberate thorax adaptation and the evolution of Grylloblattodea.
Few entomologists have e'er collected these unique insects, and little is known about their life history and biology. Withal, the potential distribution areas of the world are relative broad according our prediction in this study. Industrial development, man activities and global warming may threaten unknown and undiscovered grylloblattids. Several conservation comments are also provided.
Results
Taxonomy
Genus Grylloblattella Storozhenko, 1988 [xi]
Diagnosis: Grylloblattella can be distinguished from other genus in Grylloblattidae as follows: eyes blackness; antennae 27–38-segmented, epicranial suture not reaching the circumantennal suture; lacinia with ane or two teeth; posterior margin of pronotum incurved, without marginal expanse; tarsal pulvilli visible; cerci 9–ten-segmented; supra-anal plate of male person symmetrical, project on the posterior margin with broadly rounded or truncate tip.
Grylloblattella cheni Bai, Wang et Yang sp. nov. (Figs 1A, 2A–I)
Grylloblattella cheni Bai, Wang et Yang sp. nov., female.
(A) Habitus, lateral view. (B) Head, dorsal view. (C) Pronotum, dorsal view. (D) Cervical sclerites and eusterna of prothorax, ventral view. (Eastward) Ovipositor, lateral view. (F) Ovipositor, dorsal view. (1000) Basal antennomeres, left. (H) Lacinia with two preapical teeth, left. (I) Cercus, left.
urn:lsid:zoobank.org:act:E0375431-6D0E-4949-B0A0-EA99A5927104
Holotype: Female, Red china, Xinjiang Province, Buerjin County, Kanas Nature Reserve, viii km west to Akekule Lake (White Lake), north of Kanas Lake, south-east of Mt. Youyifeng (Friendship Elevation), N49.04173°, E87.49166°, 1750 m, raining, 2009-Vii-24; collected by Ke-Qing SONG; deposited in the collection of the Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, People's Democracy of China.
Etymology: This species is named in award of Prof. Sicien Chen (Shixiang Chen), Young man of Chinese University of Sciences, old PI for the Group of Morphology and Development of Coleoptera, Found of Zoology, Chinese Academy of Sciences, Beijing, Cathay. Prof. Chen was the founder and former manager of the Establish of Zoology, Chinese Academy of Sciences, and he fabricated swell contributions to entomological research of China.
Diagnosis: This new species tin can be attribute to Grylloblattella as follows: eyes blackness, epicranial suture not reaching the circumantennal suture, posterior margin of pronotum incurved, and tarsal pulvilli visible. Additionally, it tin can be distinguished from other species in Grylloblattidae as follows: antennae 38-segmented, cervical sclerites with five setae on each of the two lateral margins, lacinia with two teeth, and cerci 10-segmented. The key to species of Grylloblattella is given (Tabular array ane).
Tabular array 1
Key to species of Grylloblattella.
| 1 | Antennae 27–29-segmented, cerci 9-segmented, not attaining the distal stop of the 7th cercomere…………………………………..……….……2 |
| i′ | Antennae 38-segmented, cerci 10-segmented, not attaining the distal end of the 6th cercomere……………G. cheni Bai, Wang et Yang sp. nov. |
| ii(i) | Cervical sclerites with two setae on each of the ii lateral margins, basisternum oviform, cerci six.2–6.three mm in length…..…………G. sayanensis Storozhenko, 1996 [12] |
| 2′ | cervical sclerites with five setae on each of the two lateral margins, basisternum triangular, cerci 4.9 mm in length………Chiliad. pravdini (Storozhenko et Oliger, 1984) [13] |
Description: Female (holotype). Full body length 14.0 mm (measured from anterior margin of labrum to posterior margin of 10th abdominal segment) (Fig. 2A). Body colored heavy orangish-brown on head and thorax, lighter in colour on abdominal segments, and covered with numerous short hairs (Fig. 1A).
Caput attached obliquely to pronotum (Fig. 2B). Cranium wider than long (length two.2 mm, width ii.8 mm), with six setae on each lateral margin, ii setae around the antennal socket, 2 setae near eye, three on the posterior of each side and no seta on the heart; epicranial suture distinctly Y-shaped, not reaching the circumantennal suture, and a pair of parietal sutures extending from the occipital foramen over to the vertex. Clypeus two.8 times wider than long, projected on its anterior middle, with singled-out clypeal suture. Lacinia with two teeth (Fig. 2H). Eyes blackness, elongated oval in shape (Fig. 2B). Antennae filiform, composed of 38 antennomeres, the 3rd segment one.5 times as long as the 2d (Fig. 2G).
Pronotum 1.i times as long as wide, slightly concave in the posterior function, with a long median suture, some hairs on the anterior and lateral margins (Fig. 2C). Mesonotum slightly concave in the posterior office, with a long median suture. Metanotum with ii setae on its mid-lateral side and two longitudinal curt sutures in its anterior part.
Cervical sclerites about one.3 times as long as broad, triangular, elongated anteriorly, with five setae on each of the two lateral margins and minor setae on its posterior part (Fig. 2D). Basisternum of the prothorax 1.ii times as long as wide, triangular, expanded in the anterior part, with many scattered hairs. Katepisterna of the prothorax inclined, triangular in shape, situated near the posterior part of the cervical sclerite. Trochantin of the prothorax crescent shaped, with two setae on its distal part.
Legs elongate. Coxa with scattered setae and singled-out ribs on the ventral part. Profemur with one row of setae on inner margin of ventral side, no seta on lateral side of profemur; meso- and metafemur with many scattered setae. Protibia with setae on ventral side and seta on lateral side; meso- and metatibia with many scattered setae; two large spines on the apical part of all tibia. Two setae each on the apical function of the 2d and 3rd tarsi, white pulvilli and many long hairs on all tarsi, and ii potent tarsal claws without teeth. Protarsus relative length of each segment (base to apex) thirteen∶8∶7∶6∶xi. Mesotarsus relative length of each segment (base of operations to apex) xvi∶xi∶vii∶5∶ten. Metatarsus relative length of each segment (base to apex) 19∶11∶8∶4∶10.
Intestinal tergites with lateral margin flexed to the posterior, x-segmented, with ane seta each on the posterior margin of the 1st to 8th tergites, 1 seta each on the mid-lateral side of the 2nd to seventh tergites. Intestinal sternites with lateral margin flexed to the posterior, without setae.
Cercomeres 10 (length 7.2 mm), cylindrical, with one ring blueprint of setae on the distal role of all cercomeres except the first and terminal 1 (Fig. 2I). Relative length of each segment (base of operations to apex) 3∶three∶5∶8∶10∶12∶14∶16∶15∶ten. 1st cercomere without seta; 2nd cercomere with 2 setae; 3rd cercomere with iv setae; fourth cercomere with five setae; 5th cercomere with 4 setae; 6th and seventh cercomere with 4 setae each; eighth and 9th cercomere with three setae; tenth cercomere with 1 seta.
Ovipositor situated in the ventral part of the 8th and 9th intestinal segments, symmetrical, not attaining the distal end of the 6th cercomere; gonoplac (4.one mm) longer than the others, with numerous setae on the dorsal role; gonangulum situated between the 8th sternite and 1st gonapophysis, asymmetrical "/ \"-shaped, slightly pointed; 1st gonapophysis (length 3.7 mm) slightly curved medially, with numerous aptitude hairs on the lower posterior part, bordering gonangulum, and vertically emarginated mid-inductive part (Figs 2E–F).
Biological Notes: The species is known only from primary boreal coniferous forest (taiga), most Lake or river (Fig. 1B). The specimen nerveless was found under the bark of fallen dead tree, which near snow line (near two km away) in summer and could exist covered by over 1 meter deep snowfall in winter. The temperature of type locality is from 0∼x°C in summer and −xxx∼0°C in wintertime.
Geographic Distribution: This species is known from the type locality near Akekule Lake (White Lake) and north of Kanas Lake, Kanas Nature Reserve, Buerjin County, Xinjiang Province, China (Fig. 1C).
Variations of pronotum morphology in Grylloblattodea
Geometric morphometrics tin can be used to make up one's mind shape differences, and the resulting phenograms from Procrustes distances can finer indicate phenetic relationships, summarizing overall patterns of similarity [xiv]. Compared with other characters, the pronotum shows relatively loftier multifariousness in Grylloblattodea. We performed a morphometric analysis of the pronotum morphology of extant and extinct Grylloblattodea. This morphometric analysis allowed us to evaluate the similarity of the fossil pronotum to contemporary pronotum.
There were 50 equidistant semilandmarks chosen on the outline of the pronotum (tps-DIG 2.05, bend). The consensus configuration for each genus was determined and the images of each species of the genus 'unwarped' so that the semilandmarks coincided with their positions in the consensus configuration. All the species in a single genus were then superimposed onto 1 another. Analyses of the data set using tps-SMALL indicated that an splendid correlation between the tangent and the shape space existed (Fig. 3A). The correlation (uncentred) between the tangent space, Y, regressed onto Procrustes distance (geodesic distances in radians) was 0.999993. There was little doubt on the basis of the result from tps-SMALL, which supported the hypothesis that genus within a taxon such as a family tin can be analyzed by geometric morphometric methods since the results from the statistical test performed by tps-Small proved the acceptability of the data set for farther statistical analysis [14].
Pronotum shape variation exam and shape differences of 13 grylloblattids genera.
(A) Shape variation exam tps-SMALL 1.2. (B) Ordination of the 13 grylloblattids genera means along the three canonical varieties axes based on the Procrustes distance matrix. (ane) Blattogryllulus; (two) Parasheimia; (3) Plesioblattogryllus; (iv) Sojanorapbidia; (5) Sylvamicropteron; (vi) Sylvonympha; (7) Tataronympha; (8) Tillyardembia; (9) Galloisiana; (10) Grylloblatta; (xi) Grylloblattella; (12) Grylloblattina; (13) Namkungia. Green circle includes the extant 5 grylloblattids genera.
The first ii relative warps of the semilandmarks accounted for 81.85% of the variation among the genera. These were computed by a atypical-value decomposition of the weight matrix [xv]. The first two relative warps were plotted to betoken variation along the two axes (Fig. 4A). The shape changes of different genera unsaid past variation along the first two relative warp axes and shape changes were shown as deformations of the GLS reference, using thin-plate splines (Fig. 4A). The spline of each genus showed the deformation of the semilandmarks in comparison to that of the reference. The five mod genera are in the Commencement Quadrant and other eight fossil genera are in the other Quadrants.
Pronotum shape differences of 13 grylloblattids genera.
(A) Relative warps computed from the information set, plotted against 1 some other to indicate positions of the relationships amidst genera relative to one some other and to the reference configuration (situated at the origin). The shape changes of dissimilar families implied by variation along the beginning two relative warp axes. Shape changes are shown as deformations of the GLS reference, using thin-plate splines. (B) Phenetic tree (UPGMA), the trees compiled using NTSYS-pc based on Procrustes distances among the genera.
The 13 grylloblattodean genera means were plotted along the three approved varieties axes based on the Procrustes altitude matrix (Fig. 3B). UPGMA phenogram [16] of the pronotum of the 13 studied genera based on Procrustes distance matrix are presented in Figure 4B. The results signal a good correlation betwixt the scatter plots (Figs 3B, 4A) and the phenograms (produced from the Procrustes distances) (Fig. 4B). Genera clustering together in the phenograms are closely situated on the scatter plot of the first 2 relative warps.
Discussion
Phylogenetic relationships amid fossil and extant grylloblattodeans
The relationships between grylloblattodean fossil taxa and the modern grylloblattids remain unclear considering nigh fossils are based on isolated wings or have poorly preserved body features. Wheeler et al. [17] proposed that the modern Grylloblattidae were the sister group of the Dermaptera, being together the sister group of (Phasmida + Orthoptera), and that the (Plecoptera + Embioptera) were not straight related to this clade. Beutel and Gorb [xviii] claimed that the Grylloblattidae were the sister grouping of the clade {Phasmatodea + [Mantodea + (Isoptera + Blattodea)]}. Grimaldi [19] considered that the extant and fossil 'Grylloblattodea' savage in an unresolved polytomy with the majority of the other 'polyneopteran' orders. Molecular phylogenies from Terry and Whiting [20] and Cameron, Barker and Whiting [21] indicated that Grylloblattodea could be the sis grouping of the recently discovered order Mantophasmatodea, and birthday could exist the sis group of the Dictyoptera. Grimaldi and Engel [22] as well considered that the Palaeozoic and Mesozoic alate 'grylloblattids' could represent a stem group of both apterous Grylloblattodea and Mantophasmatodea. Lastly, even if Rasnitsyn [23], [24] listed several similarities between Blattogryllus and the extant Grylloblattidae, the potential synapomorphies of this last group with the fossil 'Grylloblattodea' are not articulate. We present possible relationships of all modernistic grylloblattids and eight extinct genera based on the morphology of pronotum, which is a highly diverse character in Grylloblattodea. This outcome suggests a new way to infer the phylogeny of fossil taxa and modern grylloblattids, which bridge the huge gap betwixt extremely diverse extinct winged taxa and rare extant wingless grylloblattids.
Little is known of extant grylloblattid genus and species phylogeny. Storozhenko [25] proposed a phylogeny of iv extant grylloblattid genera, Grylloblatta, Galloisiana, Grylloblattina, and Grylloblattella, based on an intuitive analysis of ten morphological and two habitat characters. A single grapheme supports the monophyly of the Asian genera (presence of 4 to eight setae on the edges of the cervical sclerites, as opposed to none in Grylloblatta), rendering the Asian genera as sister group to Grylloblatta [25]. Among the Asian genera, Storozhenko places Grylloblattina every bit sis to Galloisiana+Grylloblattella, supported by the narrow elongated condition of the right coxopodite of abdominal segment 9 of the male in Galloisiana and Grylloblattella rather than a short thickened one in Grylloblattella. Jarvis and Whiting [1] presents the kickoff-ever formal phylogenetic hypothesis of modern grylloblattid genera and species based on molecular bear witness. The topology confirms the monophyly of the three genera included in the analysis: Grylloblatta, Galloisiana, and Grylloblattina. The analysis indicates that the eastern Asian genus Galloisiana is sis to a monophyletic grouping of the east Siberian Grylloblattina and the North American Grylloblatta. Our upshot not but confirms the phylogenetic hypothesis of Grylloblatta, Galloisiana, and Grylloblattina from Jarvis and Whiting [one], but besides presents the relationships of all known modern grylloblattid genera.
Thorax evolution in Grylloblattodea
The thorax must have evolved early in the phylogenetic history of the Hexapoda as a locomotor department of the body through the specialization of its appendages for quicker motility [26]. The development of thorax morphology may be correlated with move functions involved in walking and flying.
Our results signal that there was much higher diversity in the pronotum of fossil species than in modern grylloblattids. This may be due to the totally different habitats in extant and extinct species. The food habits of the early grylloblattodeans, such every bit pollen feeding, predation, etc., were very diverse, according to the fossil record. For example, Plesioblattogryllus magnificus from Center Jurassic with the movable structures composed of the fore tarsal claws, the most apical tarsomeres, and very strong mandibles with a sharp pointed apical tooth is considered as an active hunter [27]. Winged members of Grylloblattodea might take lived in a relatively warm surroundings and a variety of habitat types in the Paleozoic and Mesozoic Era. Highly diversified pronota might reflect a various habitats and niches. Modernistic grylloblattids probably became adjusted to alive nether rocks or hidden in moss from cold areas. The lack of pronotal variation in modern grylloblattids might be explained by two hypotheses: synapomorphy or convergent evolution. The first hypothesis proposes a single clade supported by the grapheme of a virtually rectangular pronotum, which has survived afterward the extinction of other grylloblattodean taxa. The 2nd hypothesis proposes that the pronotum of the ancestors of modern grylloblattids were different in shape. After the extinction of most grylloblattodeans, the remaining species lived in similar environments, which collection convergent evolution in pronotum shape.
Insects are the only invertebrate animals which accept wings. Flying conferred an increased ability to access resources, locate mates and escape predators [28], and has undoubtedly contributed to the success of insects. Despite the obvious advantages of flight, this dispersal capacity has been lost repeatedly [29], [xxx] in near all winged orders [31]. The loss of flight, typically due to a reduction in wing length, has been attributed to the high energy expenditure required in the product and maintenance of flying apparatus, at the expense of other life-history traits [32]. Low temperature may be the central factor for the wing loss [33]. Wings are merely establish on the mesothorax and metathorax, and the prothorax never bears wings in extant insects. Mesothorax and metathorax of grylloblattids maintain very low variation in shape during ∼300 Million years evolution, which bears the wings in Paleozoic and Mesozoic Era. Nearly grylloblattid fossils contain an obviously longer meso- and metathorax than prothorax. The length of the meso- and metathorax of modern grylloblattids is unremarkably shorter than the prothorax. This may be associated with the wing loss, which is accompanied by muscle reduction and changes to the thoracic skeleton arrangement.
Threats to grylloblattids and potential distribution areas
Equally temperature is the main barriers for migration of modern grylloblattids, the distribution area of one species is very express. Migration among populations is nigh certainly severely limited or non-existent in current atmospheric condition due to grylloblattids' habitat specificity, limited geographic range of populations, and winglessness [4]. The warming of the planet since the last glaciation, compounded past human-induced global warming in recent years is causing the rapid loss of glaciers and ice sheets [34], [35], [36], [37]. In the side by side few decades, the rate at which glaciers are melting is expected to increment by two to four times from their already high rate, largely due to anthropogenic causes [34]. Grylloblattids' dependence on glacial margin habitats means that global warming is a directly threat to their futurity. Resilience of grylloblattid habitat cannot be expected without significant changes in factors linked to glacial decline.
Another potentially meaning threat to grylloblattids is development of their habitat. The known localities (Fig. 5A, black dots) of grylloblattids are very remote areas, and the potential distribution areas we inferred have environments like to those of known distribution localities. As grylloblattids can exist establish in two typical place: loftier mountain and ice cavern which might from lowland or mountain, we run the two prediction analyses. Firstly, all occurrence locations with coordinates were selected for the raw analysis, which could reflect the all possible areas (Fig. 5A). Our results evidence that the potential distribution of grylloblattid species are virtually regions populated past humans (Fig. 5A and 5B) [38]. In 2015 [39], the distribution and density of human populations be greater than in 2000 (Figs 5B–C). Human interference has acquired major ecology damage in potential and actual distribution areas of grylloblattids. Based on these evidences, we propose two areas (Fig. 5A, red circles) in which grylloblattids may possibly occur. Nosotros presume that the loftier caste of human interference in Europe (Fig. 5A, blue circle) would greatly reduce the potential to find grylloblattids there.
The comparison of the prediction map of grylloblattids and the map of Population Density of the world.
(A) Prediction of potential distribution areas of grylloblattids; black dots = selected known localities, light-green areas = potential distribution areas, red circles = the most potential areas, blueish circumvolve = the least potential areas. (B) Population Density of the globe in 2000 (after CIESIN and CIAT 2005). (C) Population Density of the world in 2015 (after CIESIN, FAO and CIAT 2005).
Secondly, simply high mountain data were used in the specific analysis for the prediction of loftier mountain grylloblattids (Fig. 6A). This map is almost same to the first prediction analysis (see Fig 5A), only a picayune bit shrinking in the prediction areas. The amplificatory map of two areas (Fig. 6A, crimson circles) indicates where grylloblattids may perhaps occur.
Prediction map of grylloblattids based on the 19 coordinates of terrestrial ice crawlers.
(A) Prediction of potential distribution areas of grylloblattids. (B) Areas in the United states of america with the well-nigh potential. (C) Areas in Asia with the about potential areas. Black dots = selected known localities, purple areas (in A) = potential distribution areas, light-green areas (in B or C) = potential distribution areas, red circles (in A) = areas with the virtually potential.
The purpose of our prediction research is non to explain the distribution of grylloblattids, simply to predict where potential areas are. The reason why many grylloblattids have been establish in Nippon (meet Fig. 5A) may be due to high homo population density close to pristine grylloblattid habitat. For instance, it is hundred-to-one that there are many more new species that could be establish in Japan because of how much piece of work has been done on grylloblattids there already. In the recent by, no new species have been establish in Japan, just in Republic of korea and China there have been several new species discovered [2], [40]. We expect that more undiscovered species could exist in many of the other predicted areas shaded green, only that it depends on 1) the accessibility of relatively undisturbed grylloblattid habitat, and 2) the number of people interested and knowledgeable enough to search for and describe new species.
The reason for the collection of Grylloblattella cheni Bai, Wang et Yang sp. nov. by Mr. Ke-Qing Song was an environmental assessment project on planned railway construction in Xinjiang, China. The report (unpublished study for North Xinjiang Environment Exploration Program) based on this assessment concluded that although the precise effects of railway construction on Grylloblattella cheni cannot be determined, in that location will certainly exist detrimental effects on this uncommonly rare species. In lodge to preserve grylloblattid habitat, we suggest the railway exist routed through lower elevation terrain, which would cause minimal disturbance to grylloblattids.
Potential distribution areas of grylloblattids are scattered over much wider areas than the very limited type localities (Fig. 5A). Virtually of these potential distribution areas are remote areas that are typically low in biodiversity. None of these areas are in the list of the well-known biodiversity hotspots, and insect surveys for conservation purposes are rarely conducted. Therefore information technology is imperative that more research be done in these regions in order to provide insight into the ecosystems that contain these unique organisms.
Materials and Methods
Nomenclatural Acts
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Geometric morphometric analysis
Digital images of grylloblattodean pronota for the morphometric analysis were from the references, edited and enhanced by Photoshop (Version nine.0). Eight fossil species belonging to eight genera and 20 extant species belonging to five genera were included in the analysis (Tabular array 2). As about fossils of Grylloblattodea are based on isolated wings or have poorly preserved body features, only eight genera with complete pronota in the fossils are involved included in the analysis (Fig. seven).
Pronotum shape of 13 grylloblattids for the geometric morphometric analysis.
(A) Blattogryllulus mongolicus Storozhenko, 1988 (Fossil). (B) Parasheimia truncata Aristov, 2004 (Fossil). (C) Plesioblattogryllus magnificus Huang, 2008 (Fossil). (D) Sojanorapbidia martynovae Storozhenko et Novokshonov, 1994 (Fossil). (E) Sylvamicropteron harpax Aristov, 2004 (Fossil). (F) Sylvonympha tshekardensis Novokshonov et Pan'kov, 1999 (Fossil). (G) Tataronympha kamensis Aristov, Novokshonov et Pan'kov, 2006 (Fossil). (H) Tillyardembia antennaeplana Zalessky, 1938 (Fossil). (I) Galloisiana chujoi Gurney, 1961. (J) Yard. kiyosawai Asahina, 1959. (K) G. kosuensis Namkung, 1974. (L) G. nipponensis (Caudell et Rex, 1924). (Grand) G. odaesanensis Kim et Lee, 2007. (N) Chiliad. olgae Vrsansky et Storozhenko, 2001. (O) K. sinensis Wang, 1987. (P) G. ussuriensis Storozhenko, 1988. (Q) G. yezoensis Asahina, 1961. (R) 1000. yuasai Asahina, 1959. (Due south) Grylloblatta barberi Caudell, 1924. (T) G. campodeiformis Walker, 1914. (U) Grand. chandleri Kamp, 1963. (V) One thousand. gurneyi Kamp, 1963. (W) M. sculleni Gurney 1937. (10) Grylloblattella cheni Bai, Wang et Yang sp. nov. (Y) G. pravdini (Storozhenko et Oliger, 1984). (Z) Grand. sayanensis Storozhenko, 1996. (AA) Grylloblattina djakonovi Bey-Bienko, 1951. (AB) Namkungia biryongensis (Namkung, 1974).
Table 2
Geometric morphometric analysis of grylloblattodean pronota represented by 8 fossil species and 20 extant species.
| Extinct | Blattogryllulus mongolicus Storozhenko, 1988 [41] |
| Parasheimia truncata Aristov, 2004 [42] | |
| Plesioblattogryllus magnificus Huang, Nel et Petrulevicius, 2008 [27] | |
| Sojanorapbidia martynovae Storozhenko et Novokshonov, 1994 [43] | |
| Sylvamicropteron harpax Aristov, 2004 [42] | |
| Sylvonympha tshekardensis Novokshonov et Pan'kov, 1999 [44] | |
| Tataronympha kamensis Aristov, Novokshonov et Pan'kov, 2006 [45] | |
| Tillyardembia antennaeplana Zalessky, 1938 [46] | |
| Extant | Galloisiana chujoi Gurney, 1961 [47] |
| G. kiyosawai Asahina, 1959 [48] | |
| G. kosuensis Namkung, 1974 [49] | |
| 1000. nipponensis (Caudell et King, 1924) [fifty] | |
| G. odaesanensis Kim et Lee, 2007 [51] | |
| K. olgae Vrsansky et Storozhenko, 2001 [2] | |
| G. sinensis Wang, 1987 [7] | |
| G. ussuriensis Storozhenko, 1988 [11] | |
| G. yezoensis Asahina, 1961 [52] | |
| G. yuasai Asahina, 1959 [48] | |
| Grylloblatta barberi Caudell, 1924 [53] | |
| G. campodeiformis Walker, 1914 [54] | |
| M. chandleri Kamp, 1963 [3] | |
| Grand. gurneyi Kamp, 1963 [three] | |
| Thousand. sculleni Gurney 1937 [55] | |
| Grylloblattella cheni Bai, Wang et Yang sp. nov. | |
| G. pravdini (Storozhenko et Oliger, 1984) [13] | |
| G. sayanensis Storozhenko, 1996 [12] | |
| Grylloblattina djakonovi Bey-Bienko, 1951 [56] | |
| Namkungia biryongensis (Namkung, 1974) [57] |
In this written report, 50 semilandmarks were selected. Photographs were input to tps-UTILS ane.38 [58] and Cartesian coordinates of semilandmarks were digitized with tps-DIG 2.05 [59]. Nosotros drew a bend along the outer margin of the pronotum outset. Then 50 semilandmarks were resampled by length for the curve. Semilandmark configurations were scaled, translated and rotated against the consensus configuration using the GLS Procrustes superimposition method [60]. Nosotros used tps-Pocket-sized 1.2 [61] to test whether the observed variation in shape was sufficiently small that the distribution of points in the tangent space can exist used as a adept approximation of their distribution in shape space (Fig. 3A). Considering shape differences betwixt genera were studied, the average or consensus configuration of semilandmarks for each genus was computed using tps-SUPER 1.14 [62]. Orthogonal to the lowest degree-squares Procrustes average configurations of semilandmarks were computed using generalized orthogonal least-squares procedures. Then the new TPS file with thirteen taxa was created for the following procedure. The coordinates were analyzed using tps-RELW ane.44 [63] to calculate eigenvalues for each principal warp (Fig. 4A). Procrustes distances (the foursquare root of the sum of squared differences between corresponding points) between each of the genera were computed and the matrix was produced by the tps-SPLIN 1.20 [64]. The Procrustes distance matrix was subjected to UPGMA (unweighted pair group method using arithmetic averages) generated by NTSYSpc [65] to determine the phenetic relationships amidst genera (Fig. 4B). The near important advantage of using Procrustes distances to capture shape variation was that these distances were considered the best method for measuring shape differences among taxa [14], [66], [67], [68], [69], [70].
Prediction of potential distribution areas of grylloblattids
Ecological niches and geographic distributional prediction of water ice crawlers were modeled using the Genetic Algorithm for Rule-ready Prediction (GARP) [71], implemented as DesktopGarp 5.1.1.vi downloaded from http://www.nhm.ku.edu/desktopgarp/Download.html. DesktopGarp is a software package for biodiversity and ecological research, which models ecological niches of species and predicts their potential distributions [71].
The geographic potential distributions were generated with GARP through a genetic algorithm that creates a series of rules matching the species specific ecological characteristics with known location occurrences [72], [73].
The species' electric current distributional points and the environmental datasets designed from groups of environmental layers were entered into DesktopGarp. The environmental data layers were bachelor through the DesktopGarp website.
The ice crawler occurrence data were obtained from the published literature, available specimen from museum or personal collections. Over hundreds of distribution data of grylloblattids were collected at start. Due to the lack of coordinates for most of them, merely 42 occurrence locations with coordinates were selected for our analysis (Table S1). As 23 occurrence locations represented water ice caves from lowland, which could non be very thought data resource for the prediction of potential distribution areas of none-cave-living water ice crawler. Nosotros ran the prediction analysis again based on the 19 coordinates of mountain ice crawler populations, for which data from ice caves were excluded.
Supporting Data
Acknowledgments
Discovery of the unique specimen resulted from Northward Xinjiang Environment Exploration Program presided by Dr. Run-Zhi Zhang, Professor of the Establish of Zoology, Chinese Academy of Sciences. We are grateful to the 2 bearding reviewers for helpful comments. Dr. Si-Qin GE, Dr. Huai-Jun XUE, and Mrs. Gan-Yan YANG from the Institute of Zoology, Chinese University of Sciences, who have given us much valuable advice on the early on version of the manuscript. We thank Mr. Jian WANG and Mr. Ji-Jiang XUE from Forestry Bureau of Aletai, who arranged the collecting trip in Kanas; Prof. Qi-Sen YANG, Dr. Lin XIA from the Institute of Zoology, Chinese Academy of Sciences and Mr. Guo-Zhong ZHANG from the Forestry Bureau of Aletai, who was very helpful with the collecting trip. Mr. Rod Crawford from Academy of Washington Burke Museum gave not bad help to Karl Jarvis for the distribution data collecting.
Footnotes
Competing Interests: The authors take alleged that no competing interests exist.
Funding: This research was partly supported past grants from the Cardinal Laboratory of Zoological Systematics and Development of the Chinese University of Sciences (No. O529YX5105), and the National Science Fund for Fostering Talents in Basic Research (Special Subjects in Animal Taxonomy, NSFC-J0630964/J0109, J0930004). Discovery of the unique specimen resulted from the North Xinjiang Environment Exploration Program presided past Dr. Run-Zhi Zhang, Professor of the Institute of Zoology, Chinese Academy of Sciences. The funders had no part in report design, data collection and assay, decision to publish, or preparation of the manuscript.
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