© CSIRO 2005							10.1071/IS04007_AC
1445-5226
Invertebrate Systematics, 2005, 19(1), 17–73.
Classification of basal Cucujoidea (Coleoptera : Polyphaga): cladistic analysis, keys and review of new families
R. A. B. LeschenA,C, J. F. LawrenceB and S. A. ŚlipińskiB
ANew Zealand Arthropod Collection, Private Bag 92170, Auckland, New Zealand.
BCSIRO Entomology, GPO Box 1700, Canberra, ACT, 2601, Australia.
CCorresponding author. Email: leschenr@landcareresearch.co.nz
Phylogenetic relationships among the basal Cucujoidea were reconstructed by a cladistic analysis of a data matrix consisting of 37 exemplar taxa and 99 adult and larval characters. Eight most parsimonious cladograms provided evidence for the polyphyly of Phloeostichidae, the paraphyly of Cucujoidea (with respect to the placement of Trogossitidae), and the monophyly of Protocucujidae + Sphindidae, Biphyllidae + Erotylidae, Cryptophagidae, Cucujidae + Silvanidae, Propalticidae + Laemophloeidae, and the Nitidulidae groups (Nitidulidae, Smicripidae, and Brachypteridae). The following families are elevated from subfamily to family status: Agapythidae (one genus), Phloeostichidae (four genera; the subfamilies Phloeostichinae and Hymaeinae are supressed), Priasilphidae (three genera), Tasmosalpingidae (one genus), and Myraboliidae (one genus). These families are described in detail and adult and larval keys to all families of Cucujoidea are provided. The genus Bunyastichus, gen. nov. (type species: B. monteithi, sp. nov.) is described in the family Phloeostichidae and the family Priasilphidae is revised with the following new taxa: Chileosilpha, gen. nov. (type species: C. elguetai, sp. nov.), Priasilpha (P. angulata, sp. nov., P. aucklandicus, sp. nov., P. bufonia, sp. nov., P. carinata, sp. nov., P. earlyi, sp. nov., and P. embersoni, sp. nov.), Priastichus (P. crowsoni, sp. nov. and P. megathorax, sp. nov.).

1. Nexus file

#NEXUS 
[MacClade 3.07 registered to Rich Leschen, Landscape Research]

BEGIN DATA;
DIMENSIONS  NTAX=37 NCHAR=99;
FORMAT SYMBOLS= " 1 2 3 4" MISSING=? GAP=- ;
CHARLABELS
	[1]	postocular_constriction	[2]	vertexal_line	[3]	median_occipital_stridulatory	[4]	paired_occipital_incisions	[5]	frontoclypeal_suture
	[6]	antennal_insertions	[7]	subantennal_groove	[8]	genal_projection	[9]	anterior_cervical_sclerites	[10]	apex_of_mandible
	[11]	dorsal_surface_of_mandible	[12]	dorsal_surface_of_mandible	[13]	mandibular_mola	[14]	galea	[15]	lacinial_uncus
	[16]	lateral_pronotal_carinae	[17]	sides_of_pronotum	[18]	anterior_pronotal_angles	[19]	anterior_portion_of_prosternum	[20]	anteromesal_corner_of_hypomero
	[21]	apex_of_prosternal_process	[22]	shortest_distance_between_proc	[23]	notosternal_suture	[24]	procoxa	[25]	protrochantin
	[26]	procoxal_cavity	[27]	procoxal_cavities_externally	[28]	procoxal_cavity	[29]	procoxal_cavity_internally	[30]	elytral_punctation
	[31]	scutellary_striole	[32]	elytral_sutural_flange	[33]	epipleuron	[34]	anteromesal_corner_of_mesepist	[35]	shortest_distance_between_meso
	[36]	'meso-metaventral_junction'	[37]	mesocoxal_cavities_laterally	[38]	metaventral_discrimen	[39]	metaventral_transverse_suture	[40]	metacoxae
	[41]	metacoxae	[42]	metacoxal_carina	[43]	metendosternal_laminae	[44]	anterior_tendons_of_metendoste	[45]	radial_cell_of_hind_wing
	[46]	free_veins_in_medial_area_of_h	[47]	tarsi_in_male	[48]	mesotarsomere_four	[49]	apex_of_terminal_tarsomere	[50]	empodium
	[51]	number_of_basal_ventrites_conn	[52]	ventrite_1	[53]	abdominal_intercoxal_process	[54]	abdominal_tergite_7	[55]	functional_spiracles_on_abdomi
	[56]	tergite_8_in_male	[57]	anterior_edge_of_sternite_8_in	[58]	apex_of_sternite_9_in_male	[59]	anterior_edge_of_sternite_9_in	[60]	base_of_tegmen
	[61]	parameres	[62]	parameres	[63]	penis	[64]	basal_portion_of_penis	[65]	base_of_penis
	[66]	anterior_edge_of_penis	[67]	'LARVA:_posterior_edge_of_head'	[68]	'LARVA:_bases_of_frontal_arms'	[69]	'LARVA:_median_endocarina'	[70]	'LARVA:_paired_endocarinae'
	[71]	'LARVA:_stemmata'	[72]	'LARVA:_antennal_length'	[73]	'LARVA:_sensorium'	[74]	'LARVA:_labrum'	[75]	'LARVA:_apex_of_mandible'
	[76]	'LARVA:_accessory_ventral_proce'	[77]	'LARVA:_mesal_surface_of_mandib'	[78]	'LARVA:_ventral_mouthparts'	[79]	'LARVA:_maxillary_articulating'	[80]	'LARVA:_inner_apical_angle_of_m'
	[81]	'LARVA:_number_of_labial_palpom'	[82]	'LARVA:_labial_palps'	[83]	'LARVA:_ligula'	[84]	'LARVA:_hypopharyngeal_sclerome'	[85]	'LARVA:_hypostomal_rods'
	[86]	'LARVA:_ventral_epicranial_ridg'	[87]	'LARVA:_gula'	[88]	'LARVA:_thoracic_and_most_abdom'	[89]	'LARVA:_mesocoxae_separated_by'	[90]	'LARVA:_number_of_pretarsal_set'
	[91]	'LARVA:_abdominal_tergites'	[92]	'LARVA:_abdominal_tergum_9'	[93]	'LARVA:_abdominal_tergum_9'	[94]	'LARVA:_urogomphi'	[95]	'LARVA:_urogomphi'
	[96]	'LARVA:_sternum_9'	[97]	'LARVA:_segment_10_and_anal_ope'	[98]	'LARVA:_spiracles'	[99]	'LARVA:_abdominal_spiracles'
	;
STATELABELS  
	1  absent present,
	2  absent present,
	3  absent present,
	4  absent present,
	5  present absent,
	6  exposed concealed,
	7  absent not_extending_below_or_behind extending_below_or_behind_eye,
	8  absent 'anterior,_truncate_or_rounded' 'anterior,_acute' lateral,
	9  not_contiguous_with_head_capsu contiguous_with_head_capsule_a apparently_absent,
	10  unidentate bidentate tridentate,
	11  without_tubercle_fitting_into with_tubercle_fitting_into_lat,
	12  without_cavity with_glabrous_cavity with_setose_cavity,
	13  present absent,
	14  at_least_2.5_X_as_wide_as_laci between_1_X_and_2.5_X_as_wide distinctly_narrower_than_lacin absent,
	15  present absent,
	16  complete incomplete_or_absent,
	17  without_4_to_6_sharp_teeth with_4_to_6_sharp_teeth,
	18  absent_or_not_produced_forward produced_forward,
	19  longer_than_prosternal_process as_long_as_as_prosternal_proce shorter_than_prosternal_proces,
	20  without_tooth with_tooth,
	21  without_lateral_projections with_lateral_projections,
	22  less_than_half_as_great_as_mid more_than_half_as_great_but_le more_than_mid_length_of_cavity,
	23  complete incomplete_or_absent,
	24  'without_or_with_short,_conceal' 'with_long,_concealed_lateral_e',
	25  exposed concealed,
	26  strongly_transverse slightly_transverse about_as_long_as_wide,
	27  open closed,
	28  without_narrow_lateral_notch 'with_narrow_lateral_notch_(les',
	29  open closed,
	30  not_seriate_or_striate seriate_or_striate,
	31  absent present,
	32  not_widened_apically widened_apically,
	33  complete_to_apex incomplete_or_absent,
	34  without_cuticular_pocket with_cuticular_pocket,
	35  less_than_half_as_great_as_sho more_than_half_as_great_but_le more_than_shortest_diameter_of,
	36  dicondylic monocondylic simple,
	37  'open_(partly_closed_by_mesepim' 'closed_(by_meeting_of_mesovent',
	38  present absent,
	39  present absent,
	40  contiguous_or_narrowly_separat widely_separated,
	41  extending_laterally_to_meet_el not_extending_laterally_to_mee,
	42  present absent,
	43  well_developed reduced absent,
	44  narrowly_separated widely_separated,
	45  'complete_(closed_basally)' incomplete_or_absent,
	46  five four three_or_fewer,
	47  '5-5-5' '5-5-4',
	48  not_or_slightly_reduced_and_no highly_reduced_and_partly_or_e,
	49  without_fringe_of_short_setae with_fringe_of_short_setae,
	50  projecting_between_pretarsal_c absent_or_not_visible_between,
	51  none two,
	52  not_much_longer_than_2 much_longer_than_2,
	53  acute_or_narrowly_rounded 'broadly_rounded,_angulate_or_t',
	54  concealed_from_above exposed_from_above,
	55  present absent,
	56  completely_dorsal with_sides_curved_ventrally with_sides_and_apex_curved_ven,
	57  without_median_strut with_median_strut,
	58  with_mesal_lobe without_mesal_lobe,
	59  without_spiculum_gastrale with_spiculum_gastrale,
	60  broadly_rounded_or_truncate narrowly_rounded_to_acute produced_anteriorly_forming_st,
	61  free_from_one_another fused_into_single_piece,
	62  articulated_to_phallobase fused_to_phallobase,
	63  not_divided_into_distinct_basa divided_into_distinct_basal_an,
	64  distinctly_wider_than_apical_p as_wide_as_or_narrower_than_ap,
	65  without_median_carina with_median_carina,
	66  without_struts with_paired_struts with_single_strut,
	67  not_or_only_slightly_emarginat distinctly_emarginate,
	68  contiguous well_separated,
	69  absent present,
	70  absent present,
	71  six five four_or_fewer,
	72  less_than_0.15_times_head_widt 0.15_to_0.5_times_head_width more_than_0.5_times_head_width,
	73  shorter_than_apical_antennomer longer_than_apical_antennomere,
	74  separated_from_head_capsule_by partly_or_completely_fused_to,
	75  unidentate bidentate tridentate,
	76  absent present,
	77  with_asperate_or_tuberculate_m with_1_to_3_hyaline_processes simple,
	78  'strongly_retracted_(distance_b' protracted_or_slightly_retract,
	79  well_developed highly_reduced_or_absent,
	80  rounded_or_truncate 'more_or_less_acute_(falciform)',
	81  two one,
	82  contiguous_or_separated_by_les separated_by_more_than_width_o,
	83  absent present,
	84  absent present,
	85  subparallel diverging absent,
	86  absent present,
	87  wider_than_long longer_than_wide,
	88  without_long_lateral_processes with_long_lateral_processes,
	89  less_than_two_coxal_diameters more_than_two_coxal_diameters,
	90  two one,
	91  without_rows_of_asperities with_curved_rows_of_asperities,
	92  not_forming_articulated_plate forming_articulated_plate,
	93  simple with_paired_urogomphi,
	94  straight curved_upwards,
	95  subparallel strongly_diverging,
	96  partly_or_entirely_exposed completely_concealed_or_appare,
	97  posterior_or_terminal posteroventral ventral,
	98  annular 'annular-biforous',
	99  not_at_ends_of_spiracular_tube at_ends_of_short_spiracular_tu at_ends_of_long_spiracular_tub at_end_of_spiracular_tubes_on,
;

MATRIX
[                                        10        20        30        40        50        60        70        80        90        ]
[                                        .         .         .         .         .         .         .         .         .         ]

Derodontus                      111121131211121121312111112112212113111111312111111111112211111112211112112211121222321111112211122
Thymalus                        111121321211121111311211111111-11122111111221211111111111223211212212121112121211111111112112211221
Eronyxa                         111222211211121111311111111111-12113111112311211111111111123111212212132112121211111112122112211321
Helota                          211122331211122112112311232122212131111221221211111121121223221212121112122211111222221111112121222
Amartus                         111212221111131111111111111121-12123111221212312121122231221111213111132111112212222211112111--1121
Australaethina                  121222321211142111312211112222112123111222312312121122231223221213121132112111212222311121112211221
Smicrips                        111111231311142111111211111121-12123122222312311121222231221111213221132112112212112112122112211221
Rhizophagus                     111222332211132111112211112122212112112122212321121212131221221212121132113211121222221111112211221
Ericmodes                       211212221311122111212211112112212112111122111121111111121223221212111122213211121222211111112221222
Protosphindus                   211111331323122121212211112112211121111122111111121111121212221212111112113211111212211111112211211
Paracucujus                     111111111323131111311111111112212113111122211212121111122122111212111113111211121222111111112111121
Chileosilpha                    111221132211121111122321112111-22121112122212211122111111122112121?????????????????????????????????
Priasilpha                      111221122211121112121311111111-22131112222111121222121111122112121121123113211121212211221112121223
Priastichus                     111221322211111112111311111111-1213312222222--21222121111122112121121133112211121212211221112111223
Hymaea                          211221142222121121111111131212222112112122112121121111111223111221221112123211121222211121212211321
Rhopalobrachium                 211221142222121211111121121211-22112112122112121122111111221111211221112123211121222211121212211321
Bunyastichus                    211212142321121121212111112112221112112122112122121111111223111211?????????????????????????????????
Phloeostichus                   211222142211121121111111111112221113112122112121121111111111111111221112123211121222211121112211321
Agapytho                        211222112211121211112111121211-22112112122212121121111111221222111211122112211121222211111112221221
Cryptamorpha                    221112112222121111111222232112212123112122222112122111111211111211111113113211121222211111122112114
Ahasverus                       211122132211111111111322232122111123112222322311122121111222121211111113113211121222211122111--2111
Cucujus                         221222132211122111111212121211-11133112112211121121111111123112221111113112212121222211121122222311
'Aulonosoma/Ancistria_first'    211222313111222112112312232112111113112122112311121111111211111212121121121132211211211122111??1222
Myrabolia                       112221232211121111112222231212121122212122212221121211111121111112211132113122121221211121112211311
Tasmosalpingus                  112121232311222112111211111121-22123122222222321121121211222121212221222213122211221212112111--13?1
Cyclaxyra                       111221312311112112311211111121-11133122122222321121111211222121213211222113122211221212121111--1221
Lamingtonium                    111221122311212112111211121212112123122122211211121111111223111211211212213122211111212121112211224
Propalticus                     221121113311111112111322221121-12133212222322311121121111221111213221231112122211??121?121121--1311
Laemophloeus                    121111131311111111112312231121-11133112222322321121121111211211213221222113122211221112122222121311
Acylomus                        111222133211111112311111231122112123212122222311121111111221221211121122213112211221212122112222111
Hobartius                       111211221221121112211211111111-12112111122212321121211111221111211111122112211122222211111112211222
Cavognatha                      111221111311121211211211221211-21233112122212111121111111213111211121112113121111211211111112111221
Atomaria                        111111131211121111111321221211-22223212222212311121221211211221221111132112211121222211111111--1111
Cryptophagus                    121221131211121112111211231111-22221212222212321121221111222221211121132112211122222211111112211221
Pharaxonotha                    111122121311121111111211231121-11222212122221211121111111111111212112122112211121222221111112211221
Xerasia                         111221111211121111311111112221-12113111122221212111111112223121212111112122111121222211112111--1121
Anchorius                       121222321211121112111111222122211122212122121211121111112123221212111112122211121222221112111--1124
;
END;

2. Eight parsimonious trees resulting from cladistic analysis

tree PAUP_1 = [&U] (1,((((2,3),36),11),((((4,(35,37)),8),(9,10)),(((((((((((((5,6),7),(25,26)),(28,29)),30),23),27),32),24),(((15,16),(17,18)),((20,21),22))),(19,(33,34))),(12,(13,14))),31))));
tree PAUP_2 = [&U] (1,((((2,3),36),11),((((4,(35,37)),(9,10)),8),(((((((((((((5,6),7),(25,26)),(28,29)),30),23),27),32),24),(((15,16),(17,18)),((20,21),22))),(19,(33,34))),(12,(13,14))),31))));
tree PAUP_3 = [&U] (1,((((2,3),36),11),((4,(9,10)),((((((((((((((((5,6),7),(25,26)),(28,29)),30),23),27),32),24),(((15,16),(17,18)),((20,21),22))),(12,(13,14))),19),(33,34)),31),8),(35,37)))));
tree PAUP_4 = [&U] (1,((((2,3),36),11),((4,(9,10)),((((((((((((((((5,6),7),(25,26)),(28,29)),30),23),27),32),24),((((15,16),17),18),((20,21),22))),(12,(13,14))),19),(33,34)),31),8),(35,37)))));
tree PAUP_5 = [&U] (1,((((2,3),36),11),((4,(9,10)),(((((((((((((((((5,6),7),(25,26)),(28,29)),30),23),27),32),24),((20,21),22)),((15,16),(17,18))),19),(12,(13,14))),31),(33,34)),8),(35,37)))));
tree PAUP_6 = [&U] (1,((2,3),((((4,(9,10)),((((((((((((((((5,6),7),(25,26)),(28,29)),30),23),27),32),24),(((15,16),(17,18)),((20,21),22))),(12,(13,14))),19),(33,34)),31),8),(35,37))),36),11)));
tree PAUP_7 = [&U] (1,((2,3),((((4,(9,10)),((((((((((((((((5,6),7),(25,26)),(28,29)),30),23),27),32),24),((((15,16),17),18),((20,21),22))),(12,(13,14))),19),(33,34)),31),8),(35,37))),36),11)));
tree PAUP_8 = [&U] (1,((2,3),((((4,(9,10)),(((((((((((((((((5,6),7),(25,26)),(28,29)),30),23),27),32),24),((20,21),22)),((15,16),(17,18))),19),(12,(13,14))),31),(33,34)),8),(35,37))),36),11)));

3. Bootstrap and jackknife trees