{"id":1024,"date":"2019-04-07T09:02:42","date_gmt":"2019-04-07T13:02:42","guid":{"rendered":"https:\/\/ibol.org\/barcodebulletin\/?p=1024"},"modified":"2020-07-16T10:54:09","modified_gmt":"2020-07-16T14:54:09","slug":"the-diversification-and-evolution-of-marine-invertebrates-in-oceanic-islands","status":"publish","type":"post","link":"https:\/\/ibol.org\/barcodebulletin\/research\/the-diversification-and-evolution-of-marine-invertebrates-in-oceanic-islands\/","title":{"rendered":"The Diversification and Evolution of Marine Invertebrates in Oceanic Islands"},"content":{"rendered":"\n[et_pb_section fb_built=”1″ fullwidth=”on” _builder_version=”3.22″ background_blend=”multiply” custom_padding=”0px|0px|0|0px|false|false”][et_pb_fullwidth_header title=”@ET-DC@eyJkeW5hbWljIjp0cnVlLCJjb250ZW50IjoicG9zdF90aXRsZSIsInNldHRpbmdzIjp7ImJlZm9yZSI6IiIsImFmdGVyIjoiIn19@” subhead=”@ET-DC@eyJkeW5hbWljIjp0cnVlLCJjb250ZW50IjoicG9zdF9leGNlcnB0Iiwic2V0dGluZ3MiOnsiYmVmb3JlIjoiIiwiYWZ0ZXIiOiIiLCJ3b3JkcyI6IiIsInJlYWRfbW9yZV9sYWJlbCI6IiJ9fQ==@” _builder_version=”3.21.4″ _dynamic_attributes=”title,subhead” content_font=”||||||||” content_text_color=”rgba(0,0,0,0.8)” subhead_font=”||on||||||” subhead_text_color=”#ffffff” subhead_font_size=”16px” background_color=”rgba(168,211,103,0)” use_background_color_gradient=”on” background_color_gradient_start=”#000000″ background_color_gradient_end=”rgba(249,195,82,0.93)” background_color_gradient_end_position=”87%” background_image=”https:\/\/ibol.org\/barcodebulletin\/wp-content\/uploads\/2019\/04\/v4.boldsystems.org_7125_animalia.png” custom_margin_tablet=”” custom_margin_phone=”” custom_margin_last_edited=”on|desktop”][\/et_pb_fullwidth_header][et_pb_fullwidth_image src=”https:\/\/ibol.org\/barcodebulletin\/wp-content\/uploads\/2019\/04\/Portugal-Figure-3.jpg” _builder_version=”3.22.2″ max_width=”95%” module_alignment=”center” custom_margin=”-20px||20px” box_shadow_style=”preset1″][\/et_pb_fullwidth_image][\/et_pb_section][et_pb_section fb_built=”1″ _builder_version=”3.22″ custom_padding=”0|0px|7px|0px|false|false”][et_pb_row module_class=” et_pb_row_fullwidth” _builder_version=”3.25″ width=”89%” width_tablet=”80%” width_phone=”” width_last_edited=”on|desktop” max_width=”89%” max_width_tablet=”80%” max_width_phone=”” max_width_last_edited=”on|desktop” custom_padding=”0|0px|27px|0px|false|false” make_fullwidth=”on”][et_pb_column type=”4_4″ _builder_version=”3.25″ custom_padding=”|||” custom_padding__hover=”|||”][et_pb_text _builder_version=”3.27.4″ text_font=”||||||||” text_text_color=”#646564″]

Macaronesian rocky shores.\nPHOTO CREDIT: Pedro Vieira, Mafalda Tavares, Henrique Queiroga<\/span>[\/et_pb_text][\/et_pb_column][\/et_pb_row][\/et_pb_section][et_pb_section fb_built=”1″ specialty=”on” _builder_version=”3.22″ custom_padding=”0|0px|54px|0px|false|false”][et_pb_column type=”3_4″ specialty_columns=”3″ _builder_version=”3.25″ custom_padding=”|||” custom_padding__hover=”|||”][et_pb_row_inner _builder_version=”3.25″][et_pb_column_inner saved_specialty_column_type=”3_4″ _builder_version=”3.25″ custom_padding=”|||” custom_padding__hover=”|||”][et_pb_text _builder_version=”3.27.4″]O<\/span>ceanic islands constitute prime evolutionary grounds for terrestrial organisms, promoting extensive isolation and harboring exceptional levels of diversity and endemism. Marine organisms, on the contrary, are not expected to experience evolutionary forces with the same intensity and, therefore, their diversification and evolution in oceanic landscapes has been somewhat disregarded.<\/p>\n

Shore-dwelling marine benthic invertebrates are unique relative to both terrestrial organisms and other marine taxa. Most shore species have planktonic larvae that facilitate dispersal over open water. However, some small invertebrates, such as free-living peracarids (Peracarida: Crustacea), are more prone to isolation due to life histories characterized by direct development and putatively reduced vagility.<\/p>[\/et_pb_text][et_pb_text _builder_version=”3.27.4″ text_font=”||||||||” width=”100%”]As a result of a DNA barcode-based screening of the diversity of littoral peracarids along Macaronesia and nearby continental shores, we have found an exceptional amount of cryptic diversity, together with a very structured geographic assortment within multiple morphospecies. We investigated, with particular detail, two common and distinct peracarid taxa present in the Macaronesian archipelagos of Azores, Madeira, and Canaries, namely 3 morphospecies of the isopod genus Dynamene1<\/sup> and 7 morphospecies of the amphipod family Hyalidae2<\/sup>. We unraveled an additional 34 suspected new species (75% endemic of Macaronesia) with no apparent discriminative morphological features, including the noteworthy cases of the isopod Dynamene edwardsi<\/em> and the amphipod Apohyale stebbingi<\/em> comprising 9 and 13 putative species, respectively.\n\nAdding to this surprising cryptic diversity, lineages within each morphospecies were also frequently exclusive to one island (50%), despite the general geographic proximity of the islands within each archipelago (e.g., from as little as 50 km between Porto Santo and Madeira). Globally, the genetic divergences among lineages within a morphospecies were also comparatively high (e.g., D. edwardsi<\/em> up to 22% and A. stebbingi<\/em> up to 21%), indicating a very deep evolutionary history in the region which pre-dates the Pleistocene glacial cycles (D. edwardsi<\/em> lineages probably started diverging between 20 and 30 MYA).\n\n [\/et_pb_text][et_pb_blurb image=”https:\/\/ibol.org\/barcodebulletin\/wp-content\/uploads\/2019\/04\/Portugal-Figure-2.jpg” content_max_width=”884px” _builder_version=”3.27.4″ body_font=”||||||||” body_text_align=”right” body_text_color=”#646564″ body_font_size=”16px” custom_padding=”||50px” border_color_all=”#929292″]

Representative study species; not to scale.
PHOTO CREDIT: Pedro Vieira<\/span><\/p>[\/et_pb_blurb][et_pb_text _builder_version=”3.27.4″]

There were several noteworthy findings in these studies. First, the unexpectedly high amount of diversity and endemism in these Macaronesian marine invertebrates, even within the same archipelago. Although peracarids can be assumed to have comparatively low vagility, since they lack a planktonic larval stage, there is extensive evidence for their dispersal capability and population connectivity. Namely, both our and other authors’ studies report peracarid morphospecies displaying little genetic structure over wide geographic ranges, for example, along the European continental Atlantic coasts.<\/p>\n

Second, we were surprised to find marked geographic segregation among the newly found species. They were frequently restricted to a single island where they constituted the only representative of the cryptic complex. To a certain extent, the geographical segregation of these peracarids more closely resemble what would be expected for terrestrial organisms than marine invertebrates.<\/p>\n

 <\/p>[\/et_pb_text][et_pb_blurb image=”https:\/\/ibol.org\/barcodebulletin\/wp-content\/uploads\/2019\/04\/Portugal-Figure-1.jpg” content_max_width=”884px” _builder_version=”3.22.2″ body_font=”||||||||” body_text_align=”right” body_text_color=”#646564″ body_font_size=”16px” custom_padding=”||50px” border_color_all=”#929292″]

A. Sampling locations for each Dynamene species. B. Reduced median network of COI data from the genus Dynamene. Size of the circles are proportional to the number of similar haplotypes. Number of mutations separating each haplotype and inferred ancestors (median vectors) are displayed in black. Links displaying a single mutation do not display the number.

PHOTO CREDIT: Pedro Vieira<\/span><\/p>[\/et_pb_blurb][et_pb_text _builder_version=”3.27.4″]

Finally, the combined evidence on diversity, geographic segregation, and divergence times, unraveled long-established divergence patterns and a remarkable geographic segregation that endured over millions of years till present. Therefore, the current distribution patterns of many of these peracarids cannot be elucidated through common accounts of marine invertebrates in the north-eastern Atlantic, namely processes involving dispersal, geographic proximity or Pleistocene glacial cycles. We propose alternative mechanisms for the speciation of these invertebrates in Macaronesia, such as those involving priority effects and pre-emptive exclusion, which have been seldom evoked to explain the deep segregation in the open ocean.<\/p>\n

In the near future, we intend to investigate the genetic variability of taxa with planktonic larvae from these oceanic islands to verify if the long-term segregation patterns are exclusive of peracarid crustaceans or, instead, are more widespread patterns in marine invertebrates in Macaronesia.<\/p>[\/et_pb_text][et_pb_text disabled_on=”off|off|off” _builder_version=”3.27.4″ text_font=”||||||||” text_font_size=”16px” custom_margin=”50px||”]

References<\/h4>\n
    \n
  1. Vieira PE, Desiderato A, Holdich DM, Soares P, Creer S, Carvalho GR, Costa FO, Queiroga H (2019) Deep segregation in the open ocean: Macaronesia as an evolutionary hotspot for low dispersal marine invertebrates. Molecular Ecology<\/em>. https:\/\/doi.org\/10.1111\/mec.15052<\/a><\/li>\n
  2. Desiderato A, Costa FO, Serejo CS, Abbiati M, Queiroga H, Vieira PE (2019) Macaronesian islands as promoters of diversification in amphipods: The remarkable case of the family Hyalidae (Crustacea, Amphipoda). Zoologica Scripta. <\/em>https:\/\/doi.org\/10.1111\/zsc.12339<\/a><\/li>\n<\/ol>[\/et_pb_text][\/et_pb_column_inner][\/et_pb_row_inner][\/et_pb_column][et_pb_column type=”1_4″ _builder_version=”3.25″ custom_padding=”|||” custom_padding__hover=”|||”][et_pb_text _builder_version=”3.27.4″ text_font=”||on||||||” text_text_color=”#646564″ text_orientation=”center”]

    Written by<\/p>[\/et_pb_text][et_pb_team_member name=”Pedro Vieira” position=”University of Minho, Braga, Portugal” image_url=”https:\/\/ibol.org\/barcodebulletin\/wp-content\/uploads\/2019\/04\/Pedro_Vieira.png” _builder_version=”3.21.4″ header_font=”||||||||” header_text_align=”center” body_font=”||||||||” text_orientation=”center” border_radii_image=”on|50px|50px|50px|50px”][\/et_pb_team_member][et_pb_team_member name=”Filipe Costa” position=”University of Minho, Braga, Portugal” image_url=”https:\/\/ibol.org\/barcodebulletin\/wp-content\/uploads\/2019\/04\/Filipe_Costa.png” _builder_version=”3.22.2″ header_font=”||||||||” header_text_align=”center” header_font_size=”18px” body_font=”||||||||” body_font_size=”14px” text_orientation=”center”][\/et_pb_team_member][et_pb_divider _builder_version=”3.21.4″ custom_margin=”||1px”][\/et_pb_divider][et_pb_text _builder_version=”3.27.4″ _dynamic_attributes=”content” text_font=”||||||||” text_text_color=”#003254″ text_orientation=”center”]@ET-DC@eyJkeW5hbWljIjp0cnVlLCJjb250ZW50IjoicG9zdF9kYXRlIiwic2V0dGluZ3MiOnsiYmVmb3JlIjoiIiwiYWZ0ZXIiOiIiLCJkYXRlX2Zvcm1hdCI6ImRlZmF1bHQiLCJjdXN0b21fZGF0ZV9mb3JtYXQiOiIifX0=@[\/et_pb_text][et_pb_button button_url=”https:\/\/ibol.org\/barcodebulletin\/wp-content\/uploads\/2019\/09\/2019_Vieira.pdf” url_new_window=”on” button_text=”Download PDF” button_alignment=”center” _builder_version=”4.4.7″ custom_button=”on” button_text_size=”12px” button_text_color=”#ffffff” button_bg_color=”#003254″ button_icon=”%%26%%” button_icon_color=”#ffffff” hover_enabled=”0″ box_shadow_style=”preset3″ box_shadow_horizontal=”3px” box_shadow_vertical=”11px”][\/et_pb_button][et_pb_text _builder_version=”4.4.7″ text_font_size=”13px” text_orientation=”center” custom_margin=”|||-36px|false|false” custom_margin_tablet=”|||0px|false|false” custom_margin_phone=”” custom_margin_last_edited=”on|tablet” hover_enabled=”0″]

    doi: 10.21083\/ibol.v9i1.5477<\/pre>\n[\/et_pb_text][et_pb_divider _builder_version=”3.21.4″][\/et_pb_divider][et_pb_sidebar area=”et_pb_widget_area_1″ show_border=”off” _builder_version=”3.21.4″][\/et_pb_sidebar][\/et_pb_column][\/et_pb_section][et_pb_section fb_built=”1″ _builder_version=”3.22″ background_color=”#000000″ custom_padding=”0|0px|2px|0px|false|false”][et_pb_row module_class=” et_pb_row_fullwidth” _builder_version=”3.25″ width=”89%” width_tablet=”80%” width_phone=”” width_last_edited=”on|desktop” max_width=”89%” max_width_tablet=”80%” max_width_phone=”” max_width_last_edited=”on|desktop” make_fullwidth=”on”][et_pb_column type=”4_4″ _builder_version=”3.25″ custom_padding=”|||” custom_padding__hover=”|||”][et_pb_post_nav in_same_term=”on” _builder_version=”3.21.4″ title_font=”|700||on|||||” title_text_color=”#ffffff”][\/et_pb_post_nav][\/et_pb_column][\/et_pb_row][\/et_pb_section][et_pb_section fb_built=”1″ disabled_on=”off|off|off” admin_label=”Follow Along” module_id=”subscribe” _builder_version=”3.29.3″ background_color=”#f7f7f7″ use_background_color_gradient=”on” background_color_gradient_start=”#d8d8d8″ background_color_gradient_end=”rgba(255,255,255,0)” background_color_gradient_end_position=”52%” background_color_gradient_overlays_image=”on” background_image=”https:\/\/ibol.org\/barcodebulletin\/wp-content\/uploads\/2019\/04\/v4.boldsystems.org_7125_animalia.png” parallax=”on” custom_padding=”49px||0||false|false” top_divider_height=”69px” global_module=”175″ saved_tabs=”all”][et_pb_row column_structure=”1_4,1_2,1_4″ module_class=” et_pb_row_fullwidth” _builder_version=”3.25″ width=”89%” width_tablet=”80%” width_phone=”” width_last_edited=”on|desktop” max_width=”89%” max_width_tablet=”80%” max_width_phone=”” max_width_last_edited=”on|desktop” custom_margin=”0px|||” custom_padding=”27px|0px|114px|0px|false|false” make_fullwidth=”on”][et_pb_column type=”1_4″ _builder_version=”3.25″ custom_padding=”|||” custom_padding__hover=”|||”][\/et_pb_column][et_pb_column type=”1_2″ _builder_version=”3.25″ custom_padding=”|||” custom_padding__hover=”|||”][et_pb_text _builder_version=”3.27.4″ header_font=”||||||||” header_font_size=”28px” header_2_font=”||||||||” header_2_font_size=”28px”]
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    The Barcode Bulletin moderates comments to\u00a0promote an informed and courteous conversation. Abusive, profane, self-promotional, or incoherent comments will be rejected.\u00a0<\/em><\/p>[\/et_pb_text][et_pb_comments show_avatar=”off” show_reply=”off” show_count=”off” disabled_on=”off|off|off” _builder_version=”3.29.3″ header_font=”||||||||” header_text_color=”#000000″ form_field_font=”||on||||||” form_field_text_align=”left” custom_button=”on” button_text_size=”13px” button_font=”||||||||” custom_padding=”|50px||50px|false|false”][\/et_pb_comments][\/et_pb_column][\/et_pb_row][\/et_pb_section][et_pb_section fb_built=”1″ _builder_version=”3.21.3″ custom_padding=”0|0px|0|0px|false|false” border_width_all=”-2px” border_width_top=”3px” border_color_top=”#a8d367″ global_module=”532″ saved_tabs=”all”][et_pb_row column_structure=”1_3,2_3″ make_equal=”on” _builder_version=”3.25″][et_pb_column type=”1_3″ _builder_version=”3.25″ custom_padding=”|||” custom_padding__hover=”|||”][et_pb_image src=”https:\/\/ibol.org\/barcodebulletin\/wp-content\/uploads\/2019\/10\/BarcodeBulletin.png” align_tablet=”center” align_phone=”” align_last_edited=”on|desktop” _builder_version=”3.29.3″ max_width_tablet=”38%” max_width_phone=”” max_width_last_edited=”on|phone”][\/et_pb_image][\/et_pb_column][et_pb_column type=”2_3″ _builder_version=”3.25″ custom_padding=”|||” custom_css_main_element=”margin:auto;” custom_padding__hover=”|||”][et_pb_text _builder_version=”3.27.4″ text_font=”||||||||” text_font_size=”16px”]
    About the Bulletin<\/a>\u00a0|\u00a0Glossary<\/a><\/a> | Archive<\/a> | iBOL Consortium<\/a> | iBOL News<\/a> | Contact Us<\/a><\/p>[\/et_pb_text][\/et_pb_column][\/et_pb_row][\/et_pb_section]\n","protected":false},"excerpt":{"rendered":"
    Unravelling long-established divergence patterns and remarkable geographic segregation that has endured over millions of years in marine invertebrates.<\/p>\n","protected":false},"author":13,"featured_media":1665,"comment_status":"open","ping_status":"open","sticky":false,"template":"","format":"standard","meta":{"_et_pb_use_builder":"on","_et_pb_old_content":"\n
    C. Matthews<\/em>1<\/em><\/sup>, J. Dorey<\/em>1<\/em><\/sup>, S. Groom<\/em>2<\/em><\/sup>, O. Davies<\/em>1<\/em><\/sup>, E. Freedman<\/em>1<\/em><\/sup>, J. Holder<\/em>1<\/em><\/sup>, B. Parslow<\/em>1<\/em><\/sup>, M. Schwarz<\/em>1<\/em><\/sup>, M. Stevens<\/em>3,4<\/em><\/sup><\/p>\n\n\n\n
    1 <\/sup>School of Biological Sciences, Flinders University, GPO Box 2100, Adelaide SA 5001, Australia 
    2 <\/sup>School of Agriculture, Food and Wine, University of Adelaide, SA 5005, Australia 
    3 <\/sup>South Australian Museum, GPO Box 234, Adelaide SA 5001, Australia 
    4 <\/sup>School of Pharmacy and Medical Sciences, University of South Australia, Adelaide, Australia<\/p>\n\n\n\n
    Fiji\u2019s entomological diversity has historically\nbeen considered depauperate. Recent widespread DNA barcoding efforts, however,\nfrom the South Australian Museum, Flinders University and University of South\nAustralia, along with a flurry of undergraduate, honours and PhD students, have\nhelped to uncover some of the hidden secrets of biodiversity within this\ntopographically complex archipelago.<\/p>\n\n\n\n
    Since 2010, funding from the Australian\n& Pacific Science Foundation and Australian Commonwealth New Colombo Plan, along\nwith support from students, has enabled fieldwork focused on collecting bees,\nwasps, and butterflies across all the major Fijian islands. Trekking up the\ntallest mountains, four wheel driving across challenging terrain, and following\nthe meandering rivers of inland Fiji has revealed that initial estimations of\nFiji\u2019s entomological fauna have been severely underestimated. <\/p>\n\n\n\n
    DNA barcoding over 1,000 bee specimens has increased species richness estimates from 4 species (known since 1979) up to 26 endemic species in the genus Homalictus. <\/em>Interestingly, 60% of these new species are only found above 800 m elevation which comprise a mere 2% of land area of Fiji, and often restricted to single mountain tops. From extensive barcoding, mitochondrial haplotype diversity was used to explore the level of intraspecific gene flow in the widespread species of the genus. <\/p>\n\n\n\n
    \"Haplotype<\/a>
    Haplotype network of all sequenced HOMALICTUS fijiensis (N=358) coloured by sampling locality. Hash marks represent nucleotide changes between each haplotype. Shared haplotypes represented by circles with multiple colours. Circle outline representing highland or lowland sampling. (b) Sampling map of H. fijiensis coloured by geographic sampling locality.<\/figcaption><\/figure><\/div>\n\n\n\n
    These results also indicate that gene flow is being restricted within highland localities of the most widespread Homalictus <\/em>species. Dispersal from a species home range does not appear to be occurring in Fiji, which may be presenting a contemporary model of speciation that is predominantly influenced by past climatic fluctuations. There is an estimated crown age of 400 ka for the initial Fijian Homalictus <\/em>colonisation, which would result in the genus being present for several glacial cycles. During glacial maxima, cooler climate would be ubiquitous throughout Fiji, however during glacial minima and interglacial periods there is a distinction between cool highland and warm lowland climate. DNA barcoding results indicate that the largest diversification of this genus is concordant with the most recent glacial minima, as species that were freely dispersing during glacial maxima are forced to retreat into highland refugia. Combined with the inferred haplotype networks, these results indicate that restriction due to low thermal tolerance of lowland climate is driving the extraordinary highland species richness in Fiji. Further to the work on bees, we have also started barcoding Fiji\u2019s butterfly fauna, along with the first ever species of Gasteruption<\/em>, a parasitoid wasp genus, found in Fiji. The species, Gasteruption tomanivi<\/em> (Published in Zootaxa by PhD student Ben Parslow), was found at the peak of Fiji\u2019s highest mountain. These discoveries have highlighted how little is known about the entomofauna of Fiji and how the use of DNA barcoding has helped to uncover Fiji\u2019s hidden secrets of biodiversity. <\/p>\n","_et_gb_content_width":"","_monsterinsights_skip_tracking":false,"_monsterinsights_sitenote_active":false,"_monsterinsights_sitenote_note":"","_monsterinsights_sitenote_category":0,"footnotes":""},"categories":[2],"tags":[71,72,105,24,110,67,65,102,101],"coauthors":[141,140],"class_list":["post-1024","post","type-post","status-publish","format-standard","has-post-thumbnail","hentry","category-research","tag-cryptic-species","tag-endemism","tag-evolution","tag-invertebrates","tag-islands","tag-marine","tag-portugal","tag-species-distribution","tag-species-diversity"],"_links":{"self":[{"href":"https:\/\/ibol.org\/barcodebulletin\/wp-json\/wp\/v2\/posts\/1024","targetHints":{"allow":["GET"]}}],"collection":[{"href":"https:\/\/ibol.org\/barcodebulletin\/wp-json\/wp\/v2\/posts"}],"about":[{"href":"https:\/\/ibol.org\/barcodebulletin\/wp-json\/wp\/v2\/types\/post"}],"author":[{"embeddable":true,"href":"https:\/\/ibol.org\/barcodebulletin\/wp-json\/wp\/v2\/users\/13"}],"replies":[{"embeddable":true,"href":"https:\/\/ibol.org\/barcodebulletin\/wp-json\/wp\/v2\/comments?post=1024"}],"version-history":[{"count":16,"href":"https:\/\/ibol.org\/barcodebulletin\/wp-json\/wp\/v2\/posts\/1024\/revisions"}],"predecessor-version":[{"id":4574,"href":"https:\/\/ibol.org\/barcodebulletin\/wp-json\/wp\/v2\/posts\/1024\/revisions\/4574"}],"wp:featuredmedia":[{"embeddable":true,"href":"https:\/\/ibol.org\/barcodebulletin\/wp-json\/wp\/v2\/media\/1665"}],"wp:attachment":[{"href":"https:\/\/ibol.org\/barcodebulletin\/wp-json\/wp\/v2\/media?parent=1024"}],"wp:term":[{"taxonomy":"category","embeddable":true,"href":"https:\/\/ibol.org\/barcodebulletin\/wp-json\/wp\/v2\/categories?post=1024"},{"taxonomy":"post_tag","embeddable":true,"href":"https:\/\/ibol.org\/barcodebulletin\/wp-json\/wp\/v2\/tags?post=1024"},{"taxonomy":"author","embeddable":true,"href":"https:\/\/ibol.org\/barcodebulletin\/wp-json\/wp\/v2\/coauthors?post=1024"}],"curies":[{"name":"wp","href":"https:\/\/api.w.org\/{rel}","templated":true}]}}