Alex's focal taxon

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Mantodea

Praying mantises are predatory insects belonging to the order Mantodea, which contains around 2,300 described species in 434 genera (Ehrmann, 2002). Synapomorphies characterizing Mantodea include the presence of raptorial forelegs, a well-defined postclypeus, femoral brush, lamella system, the loss of one denticle in the gizzard, and abdominal neuromeres II and III fused to the complex of neuromeres in thoracic segment III and abdominal segment I (Klass, 1998, 1999; Roy, 1999). Mantises are, along with cockroaches and termites (Blattodea and Isoptera, respectively), generally considered to be members of the Dictyoptera, but the relationships of these three groups remain controversial (Svenson & Whiting, 2004).

At present, around 100 species and 47 genera of Mantodea have been documented from Borneo, but few studies have specifically addressed the biodiversity of this order on the island (Ehrmann, 2002; Helmkampf et al., 2007).

Morphospecies Descriptions

Notes: (YS) in Tentative ID indicates that designation was suggested by mantid breeder Yen Saw at MantidForum. Time windows in Ecological Observations represent range of photographic documentation (in most cases beginning immediately after discovery of specimen). Laminated blue grid seen in some images is composed of 0.25 in. (6.34 mm) squares. Outgroup Mantis0014 was chosen to be a Metallyticus sp. because of broad consensus (Wieland, 2008) that this genus occupies a basal position within the Mantodea.

Mantis0001: Emerald terrapin mantis (Indiv202)

Tentative ID: Theopropus sp. (YS)

Note: Indiv202 is a nymph (and thus lacks developed wings); given high certainty in genus-level ID, character matrix entry for Mantis0001 was completed using digital images of Theopropus elegans.

Site(s): Niah National Park Headquarters, along boardwalk

Date: June 9, 2010

General description of morphology: ~15 mm total length. Compact form; vegetative mimic; alternating patches of light green and white, abdomen upturned

Ecological observations: Single individual found in a bush amongst exposed leaves (1:47 PM)

Behavioral observations: Slow-moving.

Mantis0002: Russet tommyknocker mantis (Indiv203)

Tentative ID: Gonypeta or Amantis sp. (YS)

Site(s): Lambir Hills

Date(s): June 13, 2010

General description of morphology: ~15 mm total length. Fairly uniform brown coloration (including wings), femur of raptorial arm shaped roughly like a wide isosceles triangle.

Ecological observations: Attracted to mercury vapor lamp with white sheet (12:10 AM - 12:11 AM)

Behavioral observations: Very rapidly moving (difficult to photograph without restraint by plastic bag). Most locomotion by walking legs, though specimen would attempt to take flight once suitably positioned.

Mantis0003: Casqued pharaoh mantis (Indiv204, Indiv211)

Tentative ID: Ceratocrania macra

Site(s): Lambir Hills National Park, Maliau Conservation Area

Date(s): 06/13/2010: 12:13-12:22; 06/27/2010: 8:20-8:24

General description of morphology: ~70 mm total length. Flexible horn-like projection from head, spindly, serrated pronotum, flanges on all walking legs

Ecological observations: Attracted to mercury vapor lamp with white sheet (12:13-12:22 AM)

Behavioral observations: Typically holds raptorial arms against pronotum and positions head so that anterior portion of body resembles a single twig. Actively attempts to escape plastic bags, even those furnished with vegetative cover, with repetitive ramming motions.

Mantis0004: Common green (Indiv205)

Tentative ID: Hierodula sp. --> Camelomantis sp. (YS)

Site(s): Lambir Hills National Park HQ

Date(s): 06/13/2010

General description of morphology: ~90 mm total length. Glossy green coloration, bulging teardrop-shaped eyes, white thorns on coxa of raptorial arms, thick, roughly cylindrical walking legs, long, pinched pronotum

Ecological observations: Attracted to mercury vapor lamp with white sheet (12:23-12:26 AM)

Behavioral observations: Positive vertical taxis. Will seize fingers in strong raptorial arms. Hesitant to move if all four walking legs are not securely gripping some surface.

Mantis0005: Red-scalped leaf mantis (Indiv206)

Tentative ID: Rhombodera sp.

Site(s): Lambir Hills National Park HQ

Date(s): 06/13/2010

General description of morphology: ~90 mm total length. Strong resemblance to live leaf (shield-like development of pronotum, glossy, dark green coloration), twin spots and pronounced ribbing on upper wings

Ecological observations: Attracted to mercury vapor lamp with white sheet (12:27-12:51 AM)

Behavioral observations: Will seize fingers in strong raptorial arms if threatened, but tolerates considerable handling. Positive vertical taxis: will take flight upon reaching vertical local maximum.

Mantis0006: Elfin hammerhead mantis (Indiv207)

Tentative ID: Tropidomantis sp.

Site(s): Lambir Hills National Park HQ; Maliau Basin Research Center

Date(s): 06/13/2010; 06/27/2010

General description of morphology: ~15 mm total length. Light green coloration, broadly-spaced eyes, long antennae.

Ecological observations: Attracted to mercury vapor lamp with white sheet (12:55-12:58 AM; 8:26-8:33 PM)

Behavioral observations: Skittish and quick to flight.

Mantis0007: Red-legged bonnethead mantis (Indiv208)

Tentative ID: Odontomantis sp. (YS)

Site(s): Pulau Gaya, Padang Point

Date(s): 06/19/2010

General description of morphology: ~15 mm total length. Dark green coloration, lower halves of walking legs reddish, bulging ovoid eyes with red tinge in lower halves

Ecological observations: Alighted on me while seated (1:30-1:33 PM).

Behavioral observations: Threads limb tips through the raptorial arms and pivots head to groom walking legs.

Mantis0008: Lacewing mantis (Indiv209)

Tentative ID: Tropidomantis sp.

Site(s): Maliau Basin Research Center

Date(s): 06/27/2010

General description of morphology: ~15 mm total length. Light green coloration, translucent wings, robust scalene femur, pale abdomen.

Ecological observations: Attracted to mercury vapor lamp with white sheet (7:22-7:26 PM)

Behavioral observations: Readily tracks finger movements with swiveling of head. Placid and generally stationary when sitting on hand, but takes flight abruptly.

Mantis0009: Bar-armed mantis (Indiv210, Indiv222)

Tentative ID: Hierodula sp.

Site(s): Maliau Basin Research Center

Date(s): 06/27/2010; 07/06/2010

General description of morphology: ~100 mm total length. Pinched pronotum, dark upper wings with twin light spots, rigid green fringe on lower wings, inner surface of raptorial arms marked with bars

Ecological observations: Attracted to mercury vapor lamp with white sheet (7:33-7:35 PM); flew into dining area illuminated with normal incandescent lights (10:37-10:49 PM)

Behavioral observations: Very aggressive: apt to seize fingers in raptorial arms and chew with mandibles (though unable to draw blood).

Mantis0010: Green-veined bark mantis (Indiv215)

Tentative ID: Theopompa sp. --> Humbertiella sp. (YS)

Site(s): Maliau Basin Research Center

Date(s): 06/27/2010

General description of morphology: ~50 mm total length. Green and black venation of wings, pronounced dorsoventral compression (almost flat side profile)

Ecological observations: Attracted to mercury vapor lamp with white sheet (8:30-8:32 PM); observed on wooden siding of elevated buildings exposed to secondary forest

Behavioral observations: Very swift-moving; struck at and successfully consumed a fly placed in the same plastic bag

Mantis0011: Crinkled leaf mantis (Indiv219)

Tentative ID: Deroplatys dessicata

Site(s): Maliau Basin Research Center

Date(s): 06/30/2010

General description of morphology: ~75 mm total length. Strong resemblance to dead leaf: flared, shield-like pronotum, brown coloration, wings crinkled at lower fringe. Surprisingly bright red inner wings.

Ecological observations: Captured outside dining area around dusk (6:14-6:26 PM)

Behavioral observations: Flight resembles that of a tumbling dead leaf: weak flutter to the ground.

Mantis0012: Green-scalped leaf mantis (Indiv220)

Tentative ID: Rhombodera sp.

Site(s): Maliau Basin Research Center

Date(s): 06/30/2010

General description of morphology: ~75 mm total length. Identical in most respects to Mantis0005 (Indiv206), but without red coloration on head and lower halves of eyes. Strong resemblance to live leaf, paired light spots on outer surface of wings, top of head and lower halves of eyes entirely green. Upper surface of abdomen bright red.

Ecological observations: Attracted to mercury vapor lamp with white sheet (6:28-6:52 PM)

Behavioral observations: Struggles with limbs if restrained by abdomen, but does not seize or strike fingers with raptorial appendages.

Mantis0013: Red-armed bark mantis (Indiv221)

Tentative ID: Theopompa sp.

Site(s): Maliau Basin Research Center

Date(s): 07/06/2010

General description of morphology: ~50 mm total length. Inner surface of raptorial arms red, pronounced dorsoventral flattening, black and white banding on walking legs, bark-like gray and white mottling of outer wings.

Ecological observations: Attracted to mercury vapor lamp with white sheet (10:37-10:49 PM).

Behavioral observations: Walking rapid and roach-like.

Mantis0014 (Outgroup)

Note: Image from Wieland (2008).

ID: Metallyticus sp. Thailand

Site(s): Pictured specimen from Thailand (genus reported from Borneo)

Date(s): N/A

General description of morphology: "Body oval and elongated, flat, of metallic colour. Prothorax hardly wider than long, (hardly reaching a quarter of the length of the abdomen), its sides almost straight and without lateral expansions, anteriorly not very narrow. Eyes strongly rounded. Head blunt, vertex flat. Anterior legs very large; the short femora sturdy; the four posterior femora simple as in more corpulent mantodeans. Abdomen caudally pointed, its apex not exactly well covered and protected by the tegmina and alae. Antennae simple." (Wieland, 2008: translation of Latin description by Westwood, 1835)

Ecological and behavioral observations: "... with their long legs they scurry along on the floor of the jungle or over the bark of trees at a great pace; the young, which are chequered with white or orange on the back, I have taken in decaying wood. These Mantids prey almost entirely on Cockroaches, and they pursue their victims with great vigour..." (Shelford, 1916)

Geographic Distribution

Key: X = documented at site (1); blank = not documented at site (0)

Morphotypes:

Genera:

Phylogeny and Discussion

Nexus File:

File:Mantis.nex

Character Matrix:






Consensus Tree from Pars Output:

File:AlexTree.png

Character-State Change in Tibia Form:

Character-State Change in Wing Arrangement:

Discussion:

Six most parsimonious trees were recovered from the Pars output of PHYLIP, perhaps underlining the necessity for additional characters (or pruning of the current character set to eliminate those most susceptible to homoplasy). Excepting the outgroup Mantis0014, the Mantodea examined in this project could be categorized into two major clades of 8 and 5 morphospecies; amongst individual characters, tibia form best distinguished these two groups. In the consensus tree, sister-species relationships united several morphospecies that superficial inspection suggested to be closely related (e.g., the two "bark mantises", Mantis0010 and Mantis0013); at the same time, the tree failed to support the monophyly of Tropidomantis (Mantis0006 and Mantis0008). These sister-species pairings tended to unite mantises of approximately equivalent size (e.g., Mantis0006 and Mantis0002), but higher-order groupings associated morphospecies of considerably variable size, perhaps testifying to the plasticity of overall length or bulk through evolutionary time.

Additional studies would ideally incorporate genetic techniques as well as morphological analysis; what's more, examination of morphological characters ought to proceed in a much more systematic fashion (e.g., taking a landmark-based geometric approach).

Works Cited

Ehrmann, R. (2002). Mantodea -- Gottesanbeterinnen der Welt [Mantodea -- Praying mantids of the world]. Natur und Tier -- Verlag GmbH, Münster.

Helmkampf, M.E., Schwarz, C.J., Beck, J. (2007). A first look at the biodiversity of praying mantids (Insecta: Mantodea) in Sabah, Borneo. Sepilok Bulletin 7, 1–13.

Klass, K.D. (1998). The proventriculus of the Dicondylia, with comments on the evolution and phylogeny in Dictyoptera and Odonata (Insecta). Zoologischer Anzeiger, 237, 15–42.

Klass, K.D. (1999). The pregenital abdomen of a mantid and a cockroach: musculature and nerve topography, with comparative remarks on other Neoptera (Insecta: Dictyoptera). Deutsche Entomologische Zeitschrift, 46, 3–42.

Roy, R. (1999). Morphology and taxonomy. The Praying Mantids (ed. by F. R. Prete, H. Wells, P. H. Wells and L. E. Hurd), pp. 19–40. The Johns Hopkins University Press, London.

Svenson, G.J., & Whiting, M.F. (2004). Phylogeny of Mantodea based on molecular data: evolution of a charismatic predator. Systematic Entomology 29, 359-370.

Wieland, F. (2008). The genus Metallyticus reviewed (Insecta: Mantodea). Species, Phylogeny and Evolution 1, 147–170.

List of Individuals

Individual Determined by Date Taxon Genus Species Morphotype
Indiv202 Alex Mantis1
Indiv202 Alex 9 June 2010 Mantis0001
Indiv202 Yen Saw 16 July 2010 Theopropus
Indiv203 Alex
Indiv203 Alex Mantis0002
Indiv203 Yen Saw 16 July 2010 Gonypeta or Amantis
Indiv204 Alex
Indiv204 Alex Mantis0003
Indiv204 Alex Ceratocrania macra
Indiv205 Yen Saw 16 July 2010 Camelomantis
Indiv205 Alex
Indiv205 Alex 16 July 2010 Hierodula
Indiv206 Alex
Indiv206 Alex 16 July 2010 Rhombodera Mantis0005
Indiv207 Alex
Indiv207 Alex 16 July 2010 Tropidomantis Mantis0006
Indiv208 Alex
Indiv208 Yen Saw 16 July 2010 Odontomantis Mantis0007
Indiv209 Alex
Indiv209 Alex 16 July 2010 Tropidomantis Mantis0008
Indiv210 Alex
Indiv210 Alex 16 July 2010 Hierodula Mantis0009
Indiv211 Alex
Indiv211 Alex 16 July 2010 Ceratocrania macra Mantis0003
Indiv212 Alex
Indiv212 Yen Saw 16 July 2010 Acromantis
Indiv213 Alex
Indiv214 Alex
Indiv214 Yen Saw 16 July 2010 Tropidomantis Mantis0006
Indiv215 Alex
Indiv215 Alex 16 July 2010 Theopompa Mantis0010
Indiv215 Yen Saw 16 July 2010 Humbertiella Mantis0010
Indiv216 Alex
Indiv217 Alex
Indiv217 Alex 16 July 2010 Hierodula
Indiv218 Alex
Indiv218 Alex 16 July 2010 Hierodula
Indiv219 Alex 16 July 2010 Deroplatys dessicata Mantis0011
Indiv219 Alex
Indiv220 Alex
Indiv220 Alex 16 July 2010 Rhombodera Mantis0012
Indiv221 Alex
Indiv221 Alex 16 July 2010 Theopompa Mantis0013
Indiv237 Alex 16 July 2010 Hierodula Mantis0009
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